Proceedings of Scotts Turfgrass Research Conference Volume 4 —Turfgrass Breeding January 1973 First row, left to right: William Daniel, John Madison, Eugene Mayer, Joseph Kuc, Lincoln Taylor, Richard Bangs. Second row, left to right: William R. Kneebone, Allen Dudeck, Leroy Nittler, William Richardson, Ming Yu, Terry Riordan. Third row, left to right: Joseph Duich, Derwyn Whalley, Jack Fisher, Joseph Higgins, John Long. Fourth row, left to right: Douglas Taylor, Reed Funk, Virgil Meier, Dale Benedict, James Gabert, Kenyon Payne. Proceedings of Scotts Turfgrass Research Conference Volume 4—Turfgrass Breeding January 1973 EDITORS: William R. Kneebone, Chief Editor Department of Agronomy and Plant Genetics University of Arizona Tucson, Arizona 85721 William H. Daniel Department of Agronomy Purdue University Lafayette, Indiana 47907 Kenyon T. Payne Department of Crop and Soil Science Michigan State University East Lansing, Michigan 48823 Richard T. Bangs William L. Schwaderer O. M. Scott & Sons Company Marysville, Ohio 43040 O. M. Scott & Sons Company • PUBLISHER A Subsidiary of ITT Marysville, Ohio 43040 Proceedings of Scotts Turfgrass Research Conference Volume 4 — Turfgrass Breeding Copyright © 1973 by the O. M. Scott & Sons Company. Printed in the United States of America. Library of Congress Catalog Card Number; 75-180289 PREFACE This volume contains the proceedings of a Turfgrass Re- search Conference on Turfgrass Breeding sponsored by and held at the 0. M. Scott § Sons Company, Marysville, Ohio on June 18-20, 1972. Workers concerned with a wide array of turfgrass species, turfgrass problems, and turfgrass condi- tions joined with members of the Scott's research staff in fruitful discussions. The formal (and some of the informal) expressions of problems and solutions are included between these covers. Four such conferences have now been held. It has been my distinct privilege to participate in two of these. I know that I speak for other participants (and for those who have been able to join later by means of published proceed- ings) in applauding 0. M. Scott § Sons for their leadership and farsighted public spirit in organizing and completing this series. I should like also to express my personal appreciation to Dick Bangs and Bill Schwaderer of Scotts and to Ken Payne and Bill Daniel, my associates in the editing process. Papers in this Proceedings may be reprinted in whole or part with the written permission of 0. M. Scott § Sons sub- ject to accurate quoting with proper credit to the authors and Scotts. William R. Kneebone Chief Editor C O N T E N TS P r e f a ce P a r t i c i p a n ts B r e e d i ng G r a s s es f or T u rf P u r p o s es C. R E ED FUNK and M O H A M ED K. A H M ED B i o c h e m i c al B a s is f or D i s e a se R e s i s t a n ce And Its P o t e n t i al A p p l i c a t i on in P l a nt B r e e d i ng J O S E PH A. KUC T e c h n i q u es f or A s s e s s i ng S e e d l i ng V i g or and E s t a b l i s h m e nt And P o t e n t i al A p p l i c a t i on in B r e e d i ng P r o g r a ms in G r a s s es R. D E R W YN B. W H A L L EY and CYRUS M. M c K E LL P e n n f i ne P e r e n n i al R y e g r a ss D e s c r i p t i on and P e r f o r m a n ce C h a r a c t e r i s t i cs J. M. DUICH and G. W. M c K EE S e e k i ng D w a r f n e ss in B l u e g r a ss W I L L I AM H. D A N I EL T he O r i e n t a t i on of t he R h i z o m es of K e n t u c ky B l u e g r a ss J O HN E. F I S H ER T u r f g r a ss R e s e a r ch in B r i t i sh C o l u m b ia D O U G L AS K. T A Y L OR T u r f g r a ss I m p r o v e m e nt at M i c h i g an S t a te K EN YON T. P A Y NE R e s e a r ch to I m p r o ve M e t h o ds of T e s t i ng V a r i e t al P u r i ty L E R OY W. N I T T L ER G e n e r al D i s c u s s i on B a s i cs in G r a ss J O HN H. MADISON I m p r o v e m e nt iii vii 1 15 33 57 73 75 91 9 5 99 107 B r e e d i ng B e h a v i or N o t es on S o me I m p o r t a nt T u r f g r a ss S p e c i es J O HN LONG, E U G E NE M A Y E R, V I R G IL M E I E R, T E R RY R I O R D A N, MING YU P l a nt V a r i e ty P r o t e c t i on Act and T u r f g r a ss B r e e d i ng J O S E PH HIGGINS B r e e d i ng S e e d ed V a r i e t i es of B e r m u d a g r a ss f or T u r f g r a ss U se W I L L I AM R. K N E E B O NE I m p r o v e m e nt of K e n t u c ky B l u e g r a ss f or t he T r a n s i t i on Z o ne L I N C O LN H. T A Y L OR T u r f g r a ss B r e e d i ng in F l o r i da A L L EN E. D U D E CK 121 135 149 155 161 P A R T I C I P A N TS D i ck B a n gs G o v e r n m e nt R e l a t i o ns M a n a g er O. M. Scott & Sons C o m p a ny M a r y s v i l l e, Ohio 43040 H a r o ld D o e l l i n g e r, D i r e c t or G o v e r n m e n t - I n d u s t ry R e l a t i o ns O. M. Scott & Sons C o m p a ny M a r y s v i l l e, Ohio 43040 W i l l i am H. D a n i el D e p a r t m e nt of A g r o n o my P u r d ue U n i v e r s i ty L a f a y e t t e, I n d i a na 479 07 A l l en E. D u d e ck A g r i c u l t u r al R e s e a r ch C e n t er U n i v e r s i ty of F l o r i da F o rt L a u d e r d a l e, F l o r i da 33314 J o s e ph M. D u i ch D e p a r t m e nt of A g r o n o my P e n n s y l v a n ia S t a te U n i v e r s i ty S t a te C o l l e g e, P e n n s y l v a n ia 16801 J o hn E. F i s h er P l a nt R e s e a r ch C a n a da D e p a r t m e nt of A g r i c u l t u re O t t a w a, O n t a r i o, C a n a da I n s t i t u te C. R e ed F u nk D e p a r t m e nt of Soils and C r o ps R u t g e rs U n i v e r s i ty N ew B r u n s w i c k, N ew J e r s ey 08903 J im G a b e rt B i o - C h e m i c al R e s e a r ch O. M. Scott & Sons C o m p a ny M a r y s v i l l e, Ohio 43040 J o s e ph H i g g i n s, E x a m i n er P l a nt V a r i e ty P r o t e c t i on O f f i ce U S D A - C o n s u m er and M a r k e t i ng S e r v i ce H y a tt s v i l l e, M a r y l a nd 20782 W i l l i am R. K n e e b o ne D e p a r t m e nt of A g r o n o my and P l a nt G e n e t i cs U n i v e r s i ty of A r i z o na T u c s o n, A r i z o na 85721 J o s e ph A. K uc D e p a r t m e nt of B i o c h e m i s t ry P u r d ue U n i v e r s i ty L a f a y e t t e, I n d i a na 47907 J o hn Long, D i r e c t or B i o - C h e m i c al R e s e a r ch O. M. Scott & Sons C o m p a ny M a r y s v i l l e, Ohio 43040 J o hn H. M a d i s on D e p a r t m e nt of E n v i r o n m e n t al H o r t i c u l t u re U n i v e r s i ty of C a l i f o r n ia D a v i s, C a l i f o r n ia 9 5616 E u g e ne M a y er B i o - C h e m i c al R e s e a r ch O. M. Scott & Sons C o m p a ny M a r y s v i l l e, Ohio 43040 L e r oy W. N i t t l e r, H e ad D e p a r t m e nt of S e ed I n v e s t i g a t i o ns N ew Y o rk S t a te A g r i c u l t u r al E x p e r i m e nt S t a t i on G e n e v a, N ew Y o rk 14456 K e n y on T. P a y ne D e p a r t m e nt of C r op and Soil S c i e n ce M i c h i g an S t a te U n i v e r s i ty E a st L a n s i n g, M i c h i g an 48823 W i l l i am R i c h a r d s on D e p a r t m e nt of A g r o n o my O k l a h o ma S t a te U n i v e r s i ty S t i l l w a t e r, O k l a h o ma 74074 T e r ry R i o r d an P r o d u ct D e v e l o p m e nt R e s e a r ch O. M. Scott & Sons C o m p a ny M a r y s v i l l e, Ohio 43040 J im S i m m o n s, D i r e c t or P r o d u ct D e v e l o p m e nt R e s e a r ch O. M. Scott & Sons C o m p a ny M a r y s v i l l e, Ohio 43040 D o u g l as K. T a y l or R e s e a r ch S t a t i on C a n a da D e p a r t m e nt of A g r i c u l t u re A g a s s i z, B. C. C a n a da L i n c o ln H. T a y l or D e p a r t m e nt of A g r o n o my V i r g i n ia P o l y t e c h n ic I n s t i t u te and S t a te U n i v e r s i ty B l a c k s b u r g, V i r g i n ia 24061 R. D e r w yn B. W h a l l ey D e p a r t m e nt of B o t a ny U n i v e r s i ty of N ew E n g l a nd A r m i d a l e, N.S.W. A u s t r a l ia 2351 M i ng Yu B i o - C h e m i c al R e s e a r ch O. M. Scott & Sons C o m p a ny M a r y s v i l l e, Ohio 43040 B R E E D I NG GRASSES F OR T U RF P U R P O S ES C. R e ed F u nk a nd M o h a m ed K. A h m ed D e p a r t m e nt of Soils and C r o ps R u t g e rs U n i v e r s i ty N ew B r u n s w i c k, N ew J e r s ey 089 03 I n t r o d u c t i on T he d e v e l o p m e nt of t r u ly o u t s t a n d i ng v a r i e t i es is one of t he i m p o r t a nt d u r a b i l i t y, and a t- g r e a t e st n e e ds of t he t u r f g r a ss i n d u s t r y. T he m o st a t t r i b u t es of t h e se v a r i e t i es s h o u ld be d e p e n d a b i l i t y, r e d u c ed e s- t a b l i s h m e nt and m a i n t e n a n ce r e q u i r e m e n t s, t r a c t i ve a p p e a r a n ce ( F u nk and P e p i n, 1971). E a ch of t h e se a t- t r i b u t es c an be h e i g h t e n ed o v er what t h ey a re n ow as we a c h i e ve i m p r o v ed g e n e t ic r e s i s t a n ce to d i s e a s e s, i n s e c t s, n e m a t o d es and w e e d s, a m o re d e s i r a b le g r o w th h a b i t, b e t t er t u r f - f o r m i ng p r o- p e r t i e s, and g r e a t er t o l e r a n ce of v a r i o us e n v i r o n m e n t al s u ch as h e at and d r o u g h t. A d a p t a t i on to s p e c i a l i z ed u s es and e n- v i r o n m e n ts is a l so i m p o r t a n t. F or e x a m p l e, b e t t er t u r f g r a s s es a re n e e d ed f or s h a d ed a r e a s, a t h l e t ic f i e l d s, c l o s e - c ut t e es and f a i r- w a y s, and f or p o or s o i l s. V a r i e t i es of u n i q ue or s t r i k i ng c o l or would be v e ry u s e f ul f or s p e c i al l a n d s c a p i ng p a t t e r ns a nd e f f e c t s. s t r e s s e s, T he l e a d i ng c o o l - s e a s on t u r f g r a ss s p e c i es i n c l u de K e n t u c ky t he b l u e g r a ss ( P oa p r a t e n s is L.), b e n t g r a s s es ( A g r o s t is s p p . ), and p e r e n n i al r y e g r a ss p e r e n ne L.). p r o v e m e nt c an and s h o u ld be m a d e. t he f i ne f e s c u es ( F e s t u ca spp.), ( L o l i um In e a ch of t h e se s p e c i es c o n s i d e r a b le g e n e t ic i m- K e n t u c ky B l u e g r a ss K e n t u c ky b l u e g r a ss is a p s e u d o g a m o u s, f a c u l t a t i ve a p o m i ct h a v i ng c o m p l ex c y t o l o g i c al c h a r a c t e r i s t i cs ( A k e r b e r g, 1942; G r a zi et a l ., 1961; T i n n e y, 1940), A l t h o u gh t he b a s ic c h r o m o s o me n u m b er of P oa is x = 7, m o st p l a n ts of P. p r a t e n s is p o s s e ss f r om 35 to 100 c h r o m o s o m e s. In f a c t, b i o t y p es r a n g i ng f r om 28 to o v er 150 c h r o m o s o m es h a ve b e en r e p o r t e d. A p o m i x i s, as it e x i s ts in K e n t u c ky b l u e g r a s s, is a v e ry a d- v a n t a g e o us m e t h od of m a i n t a i n i ng a s u p e r i or h y b r id v a r i e t y. E l i te c u l t i v a rs s u ch as M e r i o n, F y l k i ng and N u g g et m o st l i k e ly o r i g i n- a t ed as c h a n ce h y b r i ds in n a t u r e. T h ey m a i n t a in t h e ir s u p e r i or h y b r id g e n o t y pe b e c a u se n e a r ly a ll of t he s e ed is p r o d u c ed a s e x- u a l ly t h r o u gh a p o m i x is and is t h us g e n e t i c a l ly i d e n t i c al to t he m o t h er p l a n t. O t h er e l i te K e n t u c ky b l u e g r a ss p l a n ts h a ve b e en c o l l e c t ed w h i ch a re h i g h ly s e x u a l. W a r r e n 's A - 20 and W a r r e n 's A - 25 a re e x a m p l e s. H o w e v e r, t h ey m u st be p r o p a g a t ed v e g e t a- t i v e ly to m a i n t a in t h e ir s u p e r i or q u a l i t i es and v a r i e t al i d e n t i t y. A ll c u r r e n t ly a v a i l a b le b l u e g r a ss c u l t i v a rs o r i g i n a t ed as i n- d i v i d u al p l a nt s e l e c t i o ns f r om old t u rf a r e as or as a b e r r a nt p l a n ts in s e ed p r o d u c t i on f i e l ds or n u r s e ry r o ws w h e re no a t t e m pt h ad b e en m a de to c o n t r ol p a r e n t a g e. A p p r o x i m a t e ly 40% of t he p l a n ts c o l l e c t ed f r om p a s t u r e s, r o a d s i d es and o t h er old t u rf a r e as of t he N o r t h e a s t e rn U n i t ed S t a t es h a ve b e en s u f f i c i e n t ly a p o m i c t ic to m a i n t a in v a r i e t al i d e n t i ty ( F u nk and Han, 19 67). S u ch c o l l e c t i o ns w i ll c o n t i n ue to be a s o u r ce of n ew v a r i e t i es and a l so g e r m p l a sm f or h y b r i d i z a t i o n. P r e v i o us s t u d i es by D u i ch and M u s s er (19 59) a nd G r a z i, U m a e r us and A k e r b e rg (1961) h a ve d e m o n s t r a t ed t h at b l u e g r a ss c u l t i v a rs s u ch as M e r i on and F y l k i ng a re h i g h ly a p o- m i c t i c. M e r i on p r o d u c ed 3 - 4% n o n m a t e r n al p l a n t s. p r o d u c ed only 2%. F u r t h e r m o r e, m o st of t h e se a b e r r a nt p l a n ts w e re v e ry w e a k. T he m a j o r i ty of t he p l a n ts s t u d i ed by T i n n ey a nd A a m o dt (1940) and A k e r b e rg (1942) w e re a l so p r e d o m i n a n t ly a p o m i c t i c. C l a u s e n, K e ck and H i e s ey (1944-1945) w e re u n a b le to o b t a in h y b r i ds u s i ng P. p r a t e n s is as t he f e m a le p a r e nt in s p e c i f ic c r o s s es a l t h o u gh r e c i p r o c al c r o s s es w e re o f t en s u c c e s s f u l. B e c a u se of t h e se f a c t o r s, p l us t he c y t o g e n e t ic c o m p l e x i ty of K e n t u c ky b l u e g r a s s e s, an e x t e n s i ve r e s e a r ch p r o g r am h as b e en c o n d u c t ed to d e v e l op t e c h n i q u es and p r o c e d u r es r e q u i r ed to d e v e l op an e f f e c t i ve p r o g r am to i m p r o ve K e n t u c ky b l u e g r a ss by f ic h y b r i d i z a t i on ( F u nk and Han, 19 67; Han, 19 69; P e p i n, 19 P e p in and F u n k, 19 71). F y l k i ng i n t e r- i n t r a s p e c i- 71; S u b s t a n t i al n u m b e rs of b l u e g r a ss h y b r i ds h a ve b e en p r o d u c ed at R u t g e rs u s i ng a g r e e n h o u se c r o s s i ng t e c h n i q ue d e s c r i b ed by H an ( 19 69). P l a n ts g r o w i ng in f i e ld n u r s e r i es c an be b r o u g ht into t he g r e e n h o u se any t i me f r om D e c e m b er to M ay and f o r c ed to f l o w er u s i ng c o ol n i g h ts and e x t e n d ed d a y l e n g t h. S e ed of M e r i on a nd F y I k i ng p r o d u c ed in o ur g r e e n h o u se n o r m a l ly c o n t a i ns o v er 10% h y b r i d s. M a ny of t h e se h y b r i ds o r i g i n a te f r om t he f e r t i l i- z a t i on of u n r e d u c ed e g gs ( P e p in and F u n k, 19 71). T he egg of a p o- m i c t ic s t r a i ns of K e n t u c ky b l u e g r a ss n o r m a l ly i n i t i a t es d i v i s i on to f o rm t he p r o e m b r yo b e f o re or v e ry s o on a f t er t he f l o w er o p e ns ( T i n n e y, 1949). T h u s, t he f r e q u e n cy of h y b r i ds m ay be e n h a n c ed by a p p l y i ng p o l l en as e a r ly as p o s s i b l e. At N ew B r u n s w i c k, a n- t h e s is of b l u e g r a ss o c c u rs at night, n o r m a l ly b e t w e en m i d n i g ht a nd 5:00 a . m. S t u d i es by H an (19 69) and P e p in (19 71) d e m o n s t r a te t h at n o n- h y b r id a b e r r a nt p r o g e ny g e n e r a l ly s h ow s u b s t a n t i al d e p r e s s i on in n e a r ly a ll c h a r a c t e r i s t i cs r e l a t i ng to v e g e t a t i ve and r e p r o d u c t i ve v i g o r. On t he o t h er h a n d, v i g o r o us h y b r i ds c an be o b t a i n e d. A l t h o u gh t he a v e r a ge h y b r id is s o m e w h at l e ss v i g o r o us t h an t he a v e r a ge v i g or of i ts h i g h ly h e t e r o z y g o us s e l e c t ed p a r e n t s, a w o r k a b le n u m b er of h y b r i ds h a ve e x c e l l e nt v i g or and f r e- q u e n t ly s h ow p o s i t i ve t r a n s g r e s s i ve s e g r e g a t i o n. M a ny of t he m o re p r o m i s i ng F]_ h y b r i ds a re h i g h ly a p o m i c t i c. i n b r e e d i ng S e v e r al s e c o nd c y c le h y b r i ds d e v e l o p ed by u s i ng f i r st c y c le h y b r i ds as f e m a le p a r e n ts a re b e i ng e v a l u a t e d. A n u m b er of t h e se s e c o nd c y c le h y b r i ds s h ow e x c e l l e nt p e r f o r m a n ce in t u rf e v a l u a- in N ew J e r s e y. A d d i t i o n al e x p e r i e n ce w i th s u ch h y b r i ds t i on t r i a ls in c o m b i n a t i on w i th e x t e n s i ve c y t o l o g i c al s t u dy is n e e d ed to a s s e ss t he v a l ue of r e c u r r e nt b e st c u r r e nt h y b r i d s, s u ch as P - 2 7, P - 72 and P - 1 0 6, h a ve about 9 5 c h r o m o s o m e s. h i g h er c h r o m o s o me n u m b e rs w i ll be of p r a c t i c al v a l u e. It r e m a i ns to be s e en w h e t h er h y b r i ds w i th e v en i n t r a s p e c i f ic h y b r i d i z a t i o n. S o me of o ur T he c y t o g e n e t ic c o m p l e x i ty of K e n t u c ky b l u e g r a ss m a k es it n e c e s s a ry to w o rk w i th l a r ge p o p u l a t i o n s. A l s o, b e c a u se of t he s e v e re p h y s i o l o g i c al s t r e ss p l a c ed on a p l a nt by c o n t i n u ed c l o se m o w i n g, it is d i f f i c u lt or i m p o s s i b le to s e l e ct f or m a ny c h a r a c- t e r i s t i cs E x p e r i e n ce h as t h us i n d i c a t ed t he i m p o r t a n ce of s c r e e n i ng n u m b e rs of h y b r i ds and of s e l e c t i o ns in e i t h er s m a ll s e e d ed or s m a ll c l o n al p l o ts r e c e i v i ng t u rf m a i n t e n a n ce at an e a r ly s t a ge i m p o r t a nt in t u rf p e r f o r m a n ce in t he s p a c e d - p l a nt n u r s e r y. l a r ge in t he b r e e d i ng p r o g r am ( F u n k, E n g e l, and H a l i s k y, 19 67). B e c a u se of t he e x t r e me r a n ge of c o n d i t i o ns u n d er w h i ch t u rf is g r o w n, c o n- s i d e r a b le i m p r o v e m e nt n e e ds to be m a de in o ur a d v a n c ed e v a l u a t i on p r o c e d u r e s. I n t e r s p e c i f ic h y b r i d i z a t i on i n v o l v i ng t he u se of f o ur P. p r a t e n- s is v a r i e t i es as f e m a le p a r e n ts in c r o s s es w i th P oa c o m p r e s sa L. i n v o l v i ng t he h i g h ly s e x u al h as b e en s t u d i ed ( D a l e, 1971). C r o s s es v a r i e t i e s, W a r r e n 's A - 2 0, A - 25 and A - 2 6, p r o d u c ed a v e ry h i gh f r e q u e n cy of h y b r i d s. M a ny of t h e se h y b r i ds w e re v i g o r o us and p r o d u c ed c o n s i d e r a b ly m o re p a n i c l es t h an a re g e n e r a l ly f o u nd in K e n t u c ky b l u e g r a s s. H o w e v e r, a ll s u ch p l a n ts p r o d u c ed v e ry f ew or no v i a b le s e e d s. C y t o l o g i c al s t u d i es i n d i c a t ed t h at t h e se p r e- d o m i n a n t ly s t e r i le h y b r i ds o r i g i n a t ed f r om t he f e r t i l i z a t i on of d u c ed e g gs by r e d u c ed m a le g a m e t e s. T he 54 i n t e r s p e c i f ic h y b r i ds p r o d u c e d, u s i ng t he h i g h ly a p o m i c t ic B e l t u rf K e n t u c ky t he f e m a le p a r e n t, s h o w ed m o d e r a te to good s e ed f e r t i l i ty and a c- c e p t a b le s e e d l i ng v i g o r. S o me of t h e se h y b r i ds p r o d u c ed r a t h er a t- t r a c t i ve t u r f. C y t o l o g i c al s t u d i es i n d i c a t ed t h at t h e se f e r t i le h y b r i ds o r i g i n a t ed f r om t he f e r t i l i z a t i on of u n r e d u c ed e g gs by r e d u c ed m a le g a m e t e s. T h e se t r i p l o id F^ h y b r i ds a re m o d e r a t e ly to highly a p o- m i c t i c. b l u e g r a ss as r e- F i ne F e s c u es A n u m b er of p r o m i s i ng r ed f e s c ue p l a n ts h a ve b e en c o l l e c t ed l o w- t y p e s. f r om old t u rf a r e as of t he N o r t h e a s t. T h ey a re of two d i s t i n ct S o me h a ve t he n o n - c r e e p i ng or c h e w i n gs t y pe g r o w th habit s i m i l ar to J a m e s t o wn and H i g h l i g h t. T h ey p r o d u ce a f i n e, d e n s e, g r o w i n g, a t t r a c t i ve t u r f, and t he b e st l o c al s e l e c t i o ns s h ow b e t t er t u rf p e r f o r m a n ce t h an a ny of t he c o m m e r c i a l ly a v a i l a b le c u l t i v a r s. O t h er p l a n ts h a ve a s t r o ng s p r e a d i ng habit w i th r h i z o m es w h i ch c o m p a re f a v o r a b ly with o ur b e t t er b l u e g r a s s e s. T h e se s t r o n g ly s p r e a d i ng t y p es h a ve good s e e d l i ng v i g o r, a m e d i um t e x t u re and s h ow e x c e l l e nt p o t e n t i al f or u se on r o a d s i d es and in m i x t u r es with K e n t u c ky b l u e g r a ss f or l a wn t y pe t u r f. U n p u b l i s h ed d a ta f r om R u t g e rs U n i v e r s i ty s u g g e s ts t h at t he s t r o n g ly s p r e a d i ng t y pe f e s c u es a re r e p r o d u c t i v e ly i s o l a t ed f r om t he n o n - c r e e p i ng f e s c u e s. T he s p r e a d i ng t y p es s h ed p o l l en in m i d - a f t e r n o o n, w h e r e a s, r ed r ed t he c h e w i n gs t y p es s h ed p o l l en j u st b e f o re s u n r i s e. T h e se ing t y p e s, ( F e s t u ca r u b ra L. s u b s p. r u b r a ), a re r e p o r t ed to h a ve 56 c h r o m o s o m e s. T he c h e w i n gs t y p e s, ( F. r u b ra L. s u b s p. c o m m u t a ta Gaud.), h a ve 42 c h r o m o s o m e s. S o me h a rd f e s c ue ( F. l o n g i f o l ia T h u i l l ). v a r i e t i es r e c e n t ly d e v e l o p ed in E u r o pe a l so s h ow c o n s i d e r a b le p r o m i s e. s p r e a d- B e n t g r a ss B e n t g r a s s es do b e st in cool, m o i st c l i m a t e s. H o w e v e r, w h e re p r o f e s s i o n al m a n a g e m e nt c an be p r o v i d e d, it is p o s s i b le to u se t h e se g r a s s es t h r o u g h o ut a ll but t he h o t t e st r e g i o ns of t he U n i t ed S t a t e s. M a ny of t he e l i te c u l t i v a rs of c r e e p i ng b e n t g r a ss a re p r o- p a g a t ed v e g e t a t i v e ly f or u se on p u t t i ng g r e e ns and t e e s. c r e e p i ng b e n t g r a ss is g r o wn f r om Syn 1 s e e d. P e n n c r o ss B r e e d e rs of f u t u re b e n t g r a ss v a r i e t i es s h o u ld m a ke u se of s o me e x c e l l e nt c y t o g e n e t ic s t u d i es c o n d u c t ed in G r e at B r i t a in ( J o n e s, 19 56). T a b le 1. G e n o me r e l a t i o n s h i ps in A g r o s t i s* S p e c i es C h r o m o s o me N u m b er G e n o me C o n s t i t u t i on 1, A g r o s t is c a n i na L. s u b s p e c i es 14 Ai A^ c a n i na v e l v et b e n t g r a ss 2. A. c a n i na s u b s p e c i es M o n t a na H a r t m. 3. A. t e n u is Sibth. c o l o n i al b e n t g r a ss 4. A. s t o l o n i f e ra L. c r e e p i ng b e n t g r a ss 5. A. a l ba . L. R e d t op * A d a pt ed f r om J o n es ( 19 56). 28 28 28 42 Ai Ai Ai Ai A3 A 2 A3 A2 A2 A2 A3 A3 A 1 A 1 A 2 A 2 A 3 A3 H y b r i ds b e t w e en A. t e n u is and A. s t o l o n i f e ra s h o w ed low f e r- t i l i t y, w h e r e as h y b r i ds b e t w e en e i t h er A. t u n u is and A. a l ba or A. s t o l o n i f e ra and A. a l ba d e m o n s t r a t ed good f e r t i l i t y. P e r e n n i al R y e g r a ss P e r e n n i al r y e g r a ss ( L o l i um p e r e n ne L.) is a c r o s s - p o l l i n a t ed It is a c o o l - s e a s on b u n c h g r a ss s p e c i es w i th 14 c h r o m o s o m e s. b e st a d a p t ed in m a r i t i me c l i m a t es h a v i ng m i ld w i n t e rs and cool, m o i st s u m m e rs s u ch as N ew Z e a l a n d, t he B r i t i sh I s l es and t he c o a s t al a r e as of W a s h i n g t o n, O r e g on and N o r t h e rn C a l i f o r n i a. T he r y e g r a s s es a re not as w e ll a d a p t ed in c o n t i n e n t al c l i m a t es w h e re w i n t e rs a re c o ol and s u m m e rs hot. A n u m b er of n ew s y n t h e t ic v a r i e t i es of p e r e n n i al r y e g r a ss h a ve b e en s p e c i f i c a l ly d e v e l o p ed f or t u rf u s e. T h e se v a r i e t i es s h ow e x c e l l e nt a n ce f or s p o r ts t u rf in E u r o p e. In t he U n i t ed S t a t e s, t h ey p r o d u ce good t u rf on a t h l e t ic f i e l ds in t he m a r i t i me c l i m a t es of t he P a c i f ic N o r t h w e s t. T h ey a p p e ar v e ry p r o m i s i ng f or w i n t er o v e r s e e d i ng of w a rm s e a s on g r a s s es in t he South. T h r o u g h o ut m u ch of t he N o r t h- e a st t h ey a re b e i ng u s ed on s a n dy c o a s t al p l a in s o i ls w h e re K e n- t u c ky b l u e g r a ss g r o u n ds and s i m i l ar a r e as f or o v e r s e e d i ng t u rf a r e as d e v a s t a t ed by d i s e a s e, i n s e c ts or w e a r. is not w e ll a d a p t e d. T h ey a re a l so u s e f ul on s c h o ol p e r f o r m- A d d i t i o n al i m p r o v e m e n ts n e e d ed in o ur t u r f - t y pe r y e g r a ss i n c l u de b e t t er m o w i ng q u a l i t i e s, g r e a t er d i s e a se r e s i s t a n ce and b e t t er s u m m er and w i n t er p e r f o r m a n c e. A l e a f y, f i n e - t e x t u r e d, l o w - g r o w i ng t y pe w i th l e ss p e r s i s t a n ce would a l so be v e ry u s e f ul f or m i x t u r e s. R e f e r e n c es C i t ed A k e r b e r g, E. 1942. C y t o g e n e t ic s t u d i es in P oa p r a t e n s is and i ts h y b r id w i th P oa a l p i n a. H e r e d i t as 28:1-126. C l a u s e n, J ., D. D. K e c k, and W. M. H i e s e y. 1 9 4 4 - 4 5. E x p e r i m e n t al t a x o n o m y. C a r n e g ie I n s t i t u t i on of W a s h i n g t on Y e a r b o ok 4 4 : 7 1 - 8 1. D a l e, M. R. 19 71. An e v a l u a t i on of t a x o n o m ic c h a r a c t e rs and t u rf p e r f o r m a n ce of i n t e r s p e c i f ic h y b r i ds b e t w e en K e n t u c ky b l u e- g r a ss ( P oa p r a t e n s is L. ) and C a n a da b l u e g r a ss ( P oa c o m- p r e s sa L.). M. S. T h e s i s. R u t g e rs U n i v e r s i t y. D u i c h, J. M., a nd H. B. M u s s e r. 19 59. T he e x t e nt of a b e r r a n ts p r o- d u c ed by M e r i on K e n t u c ky b l u e g r a s s, P oa p r a t e n s is L. as d e- t e r m i n ed by f i r st and s e c o nd g e n e r a t i on p r o g e ny t e s t s. A g r o n. J o u r. 5 1 - 4 2 1 - 4 2 4. F u n k, C. R ., R. E. E n g e l, and P. M. H a l i s k y. 19 67. C l o n al e v a l u a- t i on of s e l e c t i o ns of K e n t u c ky b l u e g r a s s, P oa p r a t e n s is L. N ew J e r s ey A g r i c. E x p t. Sta. Bull. 8 1 8 : 5 5 - 5 6. F u n k, C. R ., and S. J. Han. 1967. R e c u r r e nt i n t r a s p e c i f ic h y b r i d i- z a t i o n: P oa p r a t e n s is L. N ew J e r s ey A g r i c. E x p t. Sta. Bull. a p r o p o s ed m e t h od of b r e e d i ng K e n t u c ky b l u e g r a s s, 8 1 8 : 3 - 1 4. F u n k, C. R ., and G. W. P e p i n. 1971. N ew d e v e l o p m e n ts in K e n t u c ky b l u e g r a s s. T he Golf S u p e r i n t e n d e nt 3 9 : 2 2 - 2 6. G r a z i, F ., M. U m a e r u s, a nd E. A k e r b e r g. 1961. O b s e r v a t i o ns of t he m o de of r e p r o d u c t i on and t he e m b r y o l o gy of P oa p r a t e n s is L. H e r e d i t as 4 7 : 4 8 0 - 5 4 1. H a n, S. J. 1969. E f f e c ts of g e n e t ic a nd e n v i r o n m e n t al f a c t o rs on A p o m i x is and t he c h a r a c t e r i s t i cs of n o n m a t e r n al p l a n ts K e n t u c ky b l u e g r a ss ( P oa p r a t e n s is L.) P h. D. U n i v e r s i t y. in T h e s i s. R u t g e rs J o n e s, K. 19 56. S p e c i es d i f f e r e n t i a t i on in A g r o s t i s. J o u r, of G e n e t i cs 5 4 : 3 7 0 - 3 9 4. P e p i n, G. W. 1971. E v a l u a t i on of s e l e c t ed t u r f, r e p r o d u c t i v e, and d i s e a se r e s p o n se c h a r a c t e r i s t i cs g e n i es of K e n t u c ky b l u e g r a ss ( P oa p r a t e n s is L.). P h . D. T h e s i s. R u t g e rs U n i v e r s i t y. in c r o s s ed and s e l f ed p r o- P e p i n, G. W., and C. R. F u n k. 1971. I n t r a s p e c i f ic h y b r i d i z a t i on as a m e t h od of b r e e d i ng K e n t u c ky b l u e g r a ss ( P oa p r a t e n s is L.) f or t u r f. C r op S c i e n ce 1 1 : 4 4 5 - 4 4 8. T i n n e y, F. W. 1940. C y t o l o gy of p a r t h e n o g e n e s is in P oa p r a t e n s i s. J o u r. A g r i c. R e s e a r ch 6 0 : 3 5 1 - 3 6 0. T i n n e y, F. W., and O. S. A a m o d t. 1940. T he p r o g e ny t e st as a m e a s u re of t he t y p es of s e ed d e v e l o p m e n t. 3 1 - 4 5 7 - 4 6 4. J o u r, of H e r e d i ty D i s c u s s i on P e r i od D r. D a n i e l: T he s t em r u st t h at h as b e en h i t t i ng N u g g e t, w e ' ve s e en in s p a ce p l a n t i n g s, p a r t i c u l a r l y, a v e ry d e f i n i te d e g r a d a t i on in s u m m e r t i me with a l m o st s e v e re l o ss in f a ll and w i n t e r. H a ve you o b s e r v ed t h is a l s o? D r. F u n k: We h a ve a v e ry d i f f i c u lt t i me k e e p i ng N u g g et a l i ve o ur s p a c e d - p l a nt n u r s e r i e s. T h e re a re t wo f a c t o rs i n v o l v ed o ur s i t u a t i o n. F i r s t, a p h i ds w e a k en t he p l a nt and, s e c o n d, r u st c a u s es a d d i t i o n al d a m a ge to t he w e a k e n ed p l a n t. If we d o n 't at l e a st c o n t r ol t he a p h i d s, we c an l o o se t he s p a c e d - p l a nt n u r s e ry of N u g g e t. V a r i e t i e s, to s o me d e g r ee w h i le o t h e r s, l i ke B e l t u r f, s h ow e x c e l l e nt s i s t a n ce to t h is c o m p l ex of a p h i ds and s t em r u s t. p l a n t ed r ow of N u g g et a d j a c e nt to a s p a ce p l a n t ed r ow of B e l t u r f, we n o t i ce a t r e m e n d o us d i f f e r e n ce in p e r f o r m a n c e. l i ke A n h e u s e r, s h ow t h is d a m a ge r e- In a s p a ce in in s t em D r. P a y n e: In t he P oa c o m p r e s sa h y b r i d s, ^ire you looking at t h em f r om g e n e t ic i n t e r e st or do you h o pe to t r a n s f er s o me d e- s i r a b le c h a r a c t e r i s t i cs f r om t he P. c o m p r e s s a? D r. F u n k: O ur i n i t i al i n t e r e st in m a k i ng t h is c r o ss o r i g i n a t ed f r om s o me w o rk t h at w as done in C a l i f o r n ia a n u m b er of y e a rs a go by D r. J e ns C l a u s en and his c o - w o r k e r s. T h ey f o u nd t h at it w as d i f f i c u lt or i m p o s s i b le to u se P. p r at ens is as a f e m a le p a r e nt in i n t e r s p e c i f ic h y b r i d i z a t i o n. T h ey g r ew out r a t h er l a r ge p r o g e n i es w h e re P. p r a t e n s is w as u s ed as a f e m a le p a r e n t. T h ey c o u l d n 't f i nd a ny h y b r i d s. W h e r e a s, w h en t h ey u s ed it as a m a le p a r e nt t h ey f o u nd q u i te a f ew h y b r i d s. we n o t i ce a f ew n a t u r al h y b r i ds b e t w e en P. p r a t e n s is and P. c o m p r e s s a, with P. p r a t e n s is u s ed as a f e m a le p a r e n t. We t h o u g ht w e 'd s ee if P. p r a t e n s is could be u s ed as a f e m a le p a r e nt in t h is p a r t i c u l ar c r o s s, and we f o u nd t h at o b v i o u s ly it could. T h is w as o ur i n i t i al o b j e c t i v e. p r a c t i c al s t a n d p o i nt of b r i n g i ng in g e r m p l a sm f r om P. c o m p r e s sa to i m p r o ve P. p r a t e n s is f or u se on p o or s o i l. a lot of c h a r a c t e r i s t i cs f r om P. p r a t e n s is w h i ch could be u s ed to i m p r o ve P. c o m p r e s s a. I t h i nk t h e re is s o me p o t e n t i al f r om t he I t h i nk t h e re a re J u st by c h a n ce D r. F i s h e r: Do you t h i nk t h at t h e re is a p o s s i b i l i ty t h at h i g h er c h r o m o s o me n u m b e rs m ay h a ve o c c u r r ed f r om n a t u r al h y b r i d i z a t i on in t he p a st b e- t w e en P. c o m p r e s sa and P. p r a t e n s i s? in t he a p o m i c t i cs of P oa p r a t e n s is t he i n f o r m a- c e r t a i n ly a It would j u st be s p e c u l a t i o n. T h e re is D r. F u n k: T h is is s o m e t h i ng t h at we r e a l ly h a v e n 't t he t i on to a n s w e r. p o s s i b i l i ty t h at t h is could h a ve o c c u r r e d. T he two s e em to be r e l a t ed and we k n ow t h at t he f e r t i l i z a t i on of u n r e d u c ed g a m e t es of P. p r a t e n s is by r e d u c ed g a m e t es of P. c o m p r e s sa c an give f e r t i l e, a p o m i c t ic h y b r i d s. T h e se h y b r i ds c an be r e c o g n i z ed as not t r ue P. p r a t e n s i s, but I s u s p e ct t h at if we c r o s s ed t h e se h y- b r i ds with P. p r a t e n s i s, it would be d i f f i c u lt to d i s t i n g u i sh t h e ir o f f s p r i ng f r om s o me of t he t h i n gs we h a ve b e en c a l l i ng P. p r a t e n- s i s. D r. F i s h e r: T he r e a s on I b r i ng t h is up is t h at we a re a c c u m u l a- t i ng s o me e v i d e n ce t h at N o r in 10, t he a n c e s t or of t he M e x i c an w h e a t s, h as r ye g e n es in it. We k n ow t h at N o r in 10 c a me f r om t he a n c i e nt J a p a n e se w h e a t, D a r u m a, w h i ch I am i n v e s t i g a t i ng now. D a r u ma h as r ye c h a r a c t e r i s t i cs i n t e r g e n e r ic h y b r id of h e x a p l o id w h e at and r y e. in it and m ay be an D r. Yu: C an a r e d u c ed egg of b l u e g r a ss d e v e l op into a s e ed w i t h- out f e r t i l i z a t i o n? D r. F u n k: We d o n 't h a ve good c y t o g e n e t ic d a ta on a ll of o ur c r o s s e s. H o w e v e r, we do k n ow t h at w h en we u se a h i g h ly a p o m i c t ic p l a n t, s u ch as B e l t u rf as a f e m a le p a r e n t, t h at m a ny of t he h y b r i ds r e- s u lt f r om t he f e r t i l i z a t i on of u n r e d u c ed e g g s. M a ny of t h e se h y b r i ds a re h i g h ly a p o m i c t i c. W h en we u se h i g h ly s e x u al p l a n ts of K e n t u c ky b l u e g r a s s, s u ch as W a r r e n 's A - 25 as f e m a le p a r e n t s, I s u s p e ct t h at we a re f e r t i l i z i ng m o s t ly r e d u c ed e g g s. M a ny of o ur A - 25 x A n h e u s er h y b r i ds a re h i g h ly a p o m i c t i c; w h e r e a s, m o st of o ur A - 25 x N u g g et h y b r i ds a re h i g h ly s e x u a l. D r. D. T a y l o r: You s h o w ed us p i c t u r es of c r e e p i ng f e s c u es and t he d i f f e r e nt a m o u nt of c r e e p i ng u n d er s p a ce c o n d i t i o n s. Do you o b s e r ve t he s a me d i f f e r e n c es w h en t h e y ' re c l o s e ly m o w n? Do t he c r e e p i ng t y p es f i ll in b e t t er t h an t he c h e w i n g s, or n o t? D r. F u n k: Y e s. T he old s p r e a d i ng v a r i e ty t h at we m a de a n u m- b er of y e a rs ago h as a lot of leaf spot s u s c e p t i b i l i t y, and so f o r t h, but it h as s h o wn b e t t er r e c o v e ry f r om s u m m er t h i n n i ng t h an t he c h e w i n gs t y p e s. On o ur r o a d s i de p l o t s, t he v a l ue of t he of t he s p r e a d i ng t y p es a re e v en m o re m a r k ed t h an u n d er o ur c l o s e ly m o w ed t r i a l s. W e ' ve b e en m o w i ng at t h r e e - q u a r t er in o ur f e s c ue e v a l u a t i o n s, w h i ch r e a l ly i s n 't q u i te f a ir f or s p r e a d i ng t y p e s. T h e y ' re not q u i te as t o l e r a nt of c l o se m o w i n gs as t he c h e w i n g s. I m p r o v ed t u r f - t y pe v a r i e t i es of t h e se s p r e a d i ng t y p es m ay be q u i te v a l u a b l e. r h i z o m es t h e se i n ch D r. P a y n e: You m e n t i on t he t e c h n i q ue f or g e t t i ng t he b l u e g r a s s es to s e ed in t he g r e e n h o u s e. H a ve you t r i ed t h at with t he f i ne f e s- c u es -- t he c o ol n i g h ts and long d a y s? ( A g r i c u l t u r al a f ew y e a rs ago and I w as v i s i t i ng D r. F e l ix J u s ka D r. F u n k: Y e s. R e s e a r ch S e r v i c e, USDA, B e l t s v i l l e, MD.) he m e n t i o n ed t h at he w as h a v i ng d i f f i c u l ty g e t t i ng h is f i ne f e s c u es to f l o w er in t he g r e e n h o u s e. A p p a r e n t l y, it is v e ry e a sy to c a u se f e s c u es to r e v e rt to t he v e g e t a t i ve s t a ge if you m o ve t h em to a w a rm g r e e n h o u s e. W e ' ve b e en s u c c e s s f ul in b r i n g i ng f e s c u es t he g r e e n h o u se as e a r ly as J a n u a ry if we k e ep o ur g r e e n h o u se t e m p e r a t u r es cool, e s p e c i a l ly o ur night t e m p e r a t u r e s, a nd i m m e d i a t e ly put t he p l a n ts u n d er long d a y s. T h e r e 's q u i te a d i f- f e r e n ce b e t w e en v a r i e t i es of b l u e g r a ss and f e s c u es in t h is We c an b r i ng t he c o m m on t y p es of b l u e g r a s s es h o u se w i th long d a y s, a r o u nd T h a n k s g i v i ng Day, and t h e y ' ll f l o w e r. In f a c t, we h a ve s o me v a r i e t i es of b l u e g r a ss t h at w i ll f l o w er t he g r e e n h o u s e, j u st if it is k e pt cool. t e m p e r a t u r es in t he g r e e n h o u se get down in t he 4 0 ° s, t h e y ' ll f l o w er without any o u t s i de cold t e m p e r a t u re i n d u c t i o n. O t h er v a r i e t i e s, m a ny of o ur b e t t er t u rf t y p e s, h a ve a m u ch g r e a t er w i n t er q u i r e m e n t. r e g a r d. into a cool g r e e n- If t he night into in i n c l u d i ng r e- D r. L o n g: You m e n t i o n ed t h at we h a ve a p r o b l em w i th s u m m er t h i n n i ng of b l u e g r a s s es d ue to h i gh t e m p e r a t u r e s. p h y s i o l o g i c al t y pe r e s p o n se t h at you a re s e e i n g? Is t h is a D r. F u n k: We k n ow we get m u ch p o o r er r e c o v e ry if we f e r t i l i ze h e a v i ly w i th n i t r o g e n. We k n ow t h e re a re d i s e a se p r o b l e ms i n- v o l v e d, and we k n ow t h e r e 's a t r e m e n d o us d i f f e r e n ce b e t w e en v a r i e t i es a nd t h e ir a b i l i ty to t o l e r a te t h is s u m m er c o n d i t i o n. S u m m er s u r v i v al is a c o m p l ex s i t u a t i o n. We h a ve o b s e r v ed in s u m m er v a r i e ty x y e ar and v a r i e ty x l o c a t i on i n t e r a c t i o ns l a r ge s u r v i v a l. D r. Long: What is t he m a j or d i s e a se of p e r e n n i al r y e g r a s s es y o ur r e g i o n? in D r. F u n k: In N ew J e r s e y, I would s ay t h at c r o wn r u st is p r o b a b ly t he m o st a p p a r e n t. I s l a nd and t he P a c i f ic N o r t h w e s t, p r o b a b ly C o r t i c i um r ed H e l m i n t h o s p o r i um leaf spot is a l so of s o me s i g n i f i c a n c e. In s o me of t he c o o l er a r e a s, s u ch as R h o de t h r e a d. D r. P a y n e: What t r e a t m e n ts a re you u s i ng to k e ep y o ur n u r- s e r i es f r ee of v o l u n t e e rs a nd w e e d s? D r. F u n k: We t ry to f a l l ow o ur n u r s e ry f or t wo or t h r ee m o n t hs b e f o re we t r a n s p l a nt to k i ll a ny p e r e n n i al g r a s s es and to r e d u ce t he w e ed p o p u l a t i on s o m e w h a t. T h en we a p p ly 2 , 4 -D and d i c a m ba f or w e ed c o n t r ol in t he f a l l. It d o es a good j ob on e v e r y t h i ng but a n n u al g r a s s e s, w h i ch we j u st let g r ow to give us a l i t t le w i n t er p r o t e c t i on f r om y o ur old s t e m s, and so f o r t h. T h en in t he s p r i ng we a p p ly D a c t h al at a m o d e r a t e ly h i gh r a t e. G e n e r a l l y, t h is w i ll give us p r e t ty good w e ed c o n t r ol u n t il s e ed h a r v e s t. M r. G a b e r t: What a re t he m a in d i s e a s es of t a ll f e s c ue in y o ur a r e a? D r. F u n k: T h is is s o m e t h i ng we h a v e n 't l o o k ed at. T h e r e 's c e r t a i n ly a lot of leaf s p o t t i ng and w h en w e ' ve t a k en t h em to t he p a t h o l o g i s t s, t h e y ' ve a l w a ys f o u nd t he H e l m i n t h o s p o r i um d i c t y o i d e s. T h e y ' ve a l so f o u nd m a ny o t h er o r g a n i s m s. to give an a n s w er on t h a t. O b v i o u s l y, w i n t er k i ll is a l so a p r o- b l em u n d er high n i t r o g en f e r t i l i ty and c l o se m o w i n g, e v en u n d er N ew J e r s ey c o n d i t i o n s. I j u st w o u l d n 't be a b le D r. Yu: u s ed in t a ll f e s c u e, and what t he p r o p er p r o c e d u re would b e? I would like to k n ow if hot w a t er e m a s c u l a t i on c an be D r. F u n k: We h a v e n 't a ny e x p e r i e n ce w i th hot w a t er e m a s c u l a- t i on of t a ll f e s c u e. M r. M a y e r: You r e p o r t ed t h at you t h o u g ht you s aw s o me c h l o r o s is or t ip d a m a ge on y o ur B e l t u rf and A d e l p hi b l u e g r a ss v a r i e t i es in P e n n s y l v a n ia and t h o u g ht it m i g ht be s m og i n j u r y. And t h en I b e l i e ve you t h o u g ht it m i g ht be A l t e r n a r ia or s o me o t h er d i s e a se o r g a n i s m. H a ve you c o n f i r m ed what t he p r o b l em i s? t ip D r. JFunk: t h a t. I t h i nk D r. D u i ch could give us b e t t er i n f o r m a t i on on D r. D u i c h: W e ' ve i d e n t i f i ed it as C u r v u l a r ia sp. Is t he r e a s on f or u s i ng v a r i e ty b l e n ds in b l u e- D r. L, T a y l o r: g r a ss m o re (1) t h at we d o n 't k n ow w h i ch v a r i e ty we n e ed and t h at one w i ll be d o m i n a n t, or (2) t h at b l e n ds h a ve i n t r i n s ic a d v a n- t a g es o v er s i n g le v a r i e t i e s? sp. t h at D r. D u i ch D r. F u n k: I could s ay t h at both t h i n gs a re i m p o r t a n t. W h en w e ' re r e c o m m e n d i ng v a r i e t i es f or t he g e n e r al c o n s u m er we d o n 't k n ow w h at t he s i t u a t i on w i ll be f o ur or f i ve y e a rs f r om n ow on d i s e a se c o m p l e x e s. An e x a m p le is t h is n ew C u r v u l a r ia m e n t i o n ed on B e l t u rf and s o me of t he h y b r i ds i n v o l v i ng B e l t u r f. We d o n 't k n ow t he s o i ls or m a n a g e m e nt c o n d i t i o n s. t h r ee or f o ur c o m p o n e n ts in a blend, h o p e f u l ly t he c o m p o n e nt t h a t 's b e st a d a p t ed to t he s i t u a t i on w i ll p r e d o m i n a te and give I c an s ee s o me b e t t er a d v e r se p r o b l e ms w i th b l e n ds if t h e y ' re not p r o p e r ly f o r m u l a t e d, if t h ey h a ve an o v e r ly a g g r e s s i ve c o m p o n e nt t h at is a v e ry h e a vy t h a t ch f o r m e r, or s o me o t h er p r o b l e m. I t h i nk t h at we should do m u ch m o re r e s e a r ch on b l e n d s. T he S o u t h e rn leaf blight s i t u a t i on on f i e ld c o rn i l l u s t r a t es t he i m p o r t a n ce of g e n e t ic d i v e r s i t y. A d e q u a te g e n e t ic d i v e r s i ty c an be of c o n s i d e r a b le i n s u r a n ce v a l ue in a p e r e n n i al p l a n t i n g. long t e rm d e p e n d i b i l i ty and p e r f o r m a n c e. If we h a ve D r. D a n i e l: W h en s t r i pe s m ut g e ts into a v a r i e t y, how h as a f f e c t ed s e ed p r o d u c t i on as y o u ' ve o b s e r v ed i t? it D r. F u n k: We h a v e n 't h ad a ny p r o b l em w i th s t r i pe s m ut in o ur a n n u al s e ed n u r s e r i e s. in o ur o l d er s e e d ed t u rf p l o t s, u s u a l ly t h r ee y e a rs or o l d e r. I t 's only a d i s e a se t h a t 's a p r o b l em B I O C H E M I C AL BASIS F OR D I S E A SE R E S I S T A N CE AND I TS P O T E N T I AL A P P L I C A T I ON IN P L A NT B R E E D I NG J o s e ph A. K uc D e p a r t m e nt of B i o c h e m i s t ry P u r d ue U n i v e r s i ty L a f a y e t t e, I n d i a na 47907 I n t r o d u c t i on I w i ll d i s c u ss b i o c h e m i c al m e c h a n i s ms f or d i s e a se r e- I h o pe to l e a ve you with t h r ee m e s s a g e s. One, in s i s t a n ce in p l a n t s. t h at we c an n ow b e g in to e x p l a in d i s e a se r e s i s t a n ce in p l a n ts in t e r ms of b i o c h e m i c al t e r m s - - in t e r ms of m e t a b o l ic p a t h w a y s, e n z y m e s, in t e r ms of o r g a n ic c o m p o u n d s. T he s e c o nd m e s s a ge t h at I 'd l i ke to l e a ve w i th you is t h at d i s e a se r e s i s t a n ce d o es not d e p e nd u p on t he p r e s e n ce or a b s e n ce of g e n es to do s o m e t h i n g, but w h e t h er t he g e n es a re r a p i d ly a c t i v a t ed so t h at t h ey p r o d u ce a r e s p o n se e a r ly e n o u gh and in s u f f i c i e nt m a g n i t u de to c o n t r ol t he i n f e c t i o us a g e nt w h a t e v er it m ay be -- f u n g u s, b a c t e r ia or v i r u s. t a l k i ng about t he p r e s e n ce or a b s e n ce of g e n es f or but r a t h er w h e t h er t h ey a re a c t i v a t ed e a r ly e n o u gh f i c i e nt m a g n i t u de to c o n t r ol t he i n f e c t i o us a g e n t. And f i n a l l y, t h i r d, t h at we c an m a n i p u l a te t h is g e n e t ic i n f o r m a t i o n, m u ch i m m u n i ze as a m i c r o r g a n i sm d o e s, to i n d u ce r e s i s t a n ce and to p l a n ts a g a i n st d i s e a s e, m u ch as a n i m a ls h a ve b e en i m m u n i z ed a g a i n st d i s e a s e. We w i ll s e e, in f a c t, t h at t h e re a re m a ny s i m i- l a r i t i es b e t w e en d i s e a se r e s i s t a n ce m e c h a n i s ms in p l a n ts and d i s e a se r e s i s t a n ce m e c h a n i s ms I c o n s i d er t h is a v e ry i m p o r t a nt c o n c e p t. W e ' re not r e s i s t a n c e, and in s u f- in a n i m a l s. I ' ll be s p e n d i ng m o st of my t i me d i s c u s s i ng t wo s y s t e m s. O ne is a d i s e a se of t he I r i sh p o t a t o, L a te B l i g h t, c a u s ed by t he f u n g us P h y t o p h t h o ra b u t i on to h i s t o r y; it c a u s ed t he e p i d e m ic of p o t a to in I r e l a nd w h i ch h ad s u ch d e v a s t a t i ng e f f e c ts on t he p o p u l a t i on and r e s u l t ed in m a ss m i g r a t i o n s, e s p e c i a l ly to t he U n i t ed S t a t e s. L a te B l i g ht is s t i ll a v e ry i m p o r t a nt d i s e a s e. T he o t h er d i s e a se t h at I 'm i n f e s t a n s. L a te B l i g ht m a de q u i te a c o n t r i- going to t a lk about is a d i s e a se of g r e en b e a n. T he d i s e a se b e an a n t h r a c n o s e, c a u s ed by C o l l e t o t r i c h um is l i n d e m u t h i a n u m. Why h a ve we c h o s en t h e se m o d el s y s t e ms f or o ur s t u d i e s? P o t a to t u b e rs a re e a sy to get. We h a ve g r o wn m a ny d i f f e r e nt l i n es of p o t a t o s, d i f f e r e n t i al v a r i e t i es t h at a re r e s i s t a nt to one r a ce and s u s c e p t i b le to a n o t h er of P. i n f e s t a n s. T he s a me t h i ng h o l ds t r ue f or b e a n s. We c an e a s i ly g r ow t he b e an t i s s ue and we h a ve m a i n t a i n ed m a ny c u l t i v a rs t h at d i f f er in t h e ir r e a c t i o ns to d i f f e r e nt r a c es of t he C. l i n d e m u t h i a n u m. B o th f u n gi c an be e a s i ly g r o wn in v i t r o, and d i s t i n ct r a c es of both a re a v a i l a b l e. T e s t i ng f or P o t a to D i s e a se R e s i s t a n ce P o t a to t u b e rs a re s l i c e d, put into p e t ri d i s h e s, i n o c u l a t ed a nd i n c u b a t e d. T u b er s l i c es of t he c u l t i v ar K e n n e b ec s p r a y ed with s t e r i le d e - i o n ed w a t er and i n c u b a t ed in p e t ri d i s h es f or 72 h o u rs s h ow s o me n a t u r al d i s c o l o r a t i o n. T he s a me c u l t i v ar i n o c u l a t ed w i th r a ce 124 of P. i n f e s t a ns s h o ws t he c o m p a t i b le or s u s c e p t i b le i n t e r a c t i o n. A f t er 72 h o u rs t h e re is v e ry l i t t le d i f f e r e n ce f r om t he c o n t r ol and you m i g ht be t e m p t ed to s ay t h at t h is is r e s i s t a nt t i s s u e. If you wait a n o t h er 24 h o u rs to 48 h o u r s, h o w e v e r, f u n g al g r o w th is r e a d i ly e v i d e nt on t he s u r f a ce and s p o r u l a t i on o c c u r s. T he s y m p t o ms of t he s u s c e p t i b le r e a c t i on a re not e v i d e nt f or s o me c o n s i d e r a b le t i m e, d u r i ng w h i ch t he o r g a n i sm in t he t i s s ue without a p p a r e n t ly d a m a g i ng is r a m i f y i ng it. If we p l a ce N ow l e t 's look at t he r e s i s t a nt i n t e r a c t i o n. r a ce 4 on t he c u l t i v ar K e n n e b e c, we i n d u ce a h y p e r s e n s i t i ve r e a c t i o n. T h is is an i m m e d i a te r e s p o n se t h at is e v i d e nt as e a r ly as 12 h o u rs a f t er i n o c u l a t i on and h as p r o b a b ly r e a c h ed i ts m a x i m um i n t e n s i ty a b o ut 24 h o u rs a f t er i n o c u l a t i o n. W e ' ve u s ed a h e a vy but w e ' ve a l so u s ed an e q u al i n o c u l um in i n o c u l a t i ng with t he o t h er r a ce w h e re we had a c o m p a t i b le i n t e r a c t i o n. y o u 'd be t e m p t ed to c a ll t he h y p e r s e n s i t i ve r e s p o n se to r a ce 4, t he s u s c e p t i b le i n t e r a c t i o n, and t he r e s p o n se to r a ce 124, t he r e s i s t a nt In f a c t, h y p e r s e n s i t i v i ty g i v es high t a n ce - - an e x a m p le of g r e at i n c o m p a t i b i l i t y. As s o on as o r g a n i sm g e ts into t he p l a n t, we h a ve t he d e a th of a f ew c e l ls and t he o r g a n i sm g e ts no f u r t h e r. T he o r g a n i sm is c o n t a i n e d. r e s i s- t he i n t e r a c t i o n. i n o c u l u m, In looking at t h e se r e s i s t a n c e. r e- If we i n o c u l a te t i s s ue One of t he t h o u g h ts t h at h as b e en v e ry p r e v a l e nt in d i s e a se r e- r e s i s t a n ce w o rk is t h at t h e re a re p l a n ts t h at h a ve g e n es f or s i s t a n ce and t h e re a re p l a n ts t h at d o n 't h a ve g e n es f or It h as b e en s u g g e s t ed t h at a s p e c i f ic g e ne p r o d u ct d e t e r m i n es s i s t a n ce to e a ch p a t h o g e n. T h is c o n c e pt c an be m i s l e a d i n g. We s et up s o me e x p e r i m e n ts to e x p l o re t h is c o n c e pt and to s ee w h e t h er it m i g ht be m i s l e a d i n g. We m a de t h in l a y er c h r o m a t o g r a ms of e x t r a c ts f r om t u b e rs of two c u l t i v a rs of p o t a t o, R u s s et B u r b a nk and K e n n e b e c. T he t u b e rs w e re i n o c u l a t ed w i th t wo r a c es of P. i n f e s t a ns w h i ch c an a t t a ck t he f o r m e r, but not t he l a t t er c u l t i v a r. K e n n e b ec is r e s i s t a nt and R u s s e tt B u r b a nk is s u s c e p t i b l e. We a l so i n o c u l a t ed with t he f u n g us H e l m i n t h o s p o r i um c a r b o n u m, w h i ch d o es not a t t a ck t he p o t a t o. It is a n o n - p a t h o g en of p o t a t o, but it is a p a t h o g en of c o r n. What did we s e e? t h e se t i s s u e s, e x t r a ct t h e m, r un c h r o m a t o g r a ms of t he e x t r a c t s, a nd s p r ay t he c h r o m a t o g r a ms with d i f f e r e nt r e a g e n ts w h i ch w i ll d e t e ct m e t a b o l i t e s, we s ee a p a t t e rn of m e t a b o l i t es w h i ch is a f i n g e r p r i nt f or an i n t e r a c t i o n. D e p e n d i ng u p on t he r e a g e nt we u se to s p r ay with or how we e x t r a ct t i s s u e s, we c an s ee d i f f e r e nt c o m- p o u n d s. S p r a y i ng with a n t i m o ny t r i - c h l o r i d e, f or e x a m p l e, d e t e c ts a m o ng o t h er t h i n g s, t e r p e n o i d - t y pe or s t e r o i d - t y pe c o m p o u n d s. E a ch spot on t he c h r o m a t o g r am r e p r e s e n ts at l e a st one c o m p o u nd t h at h as a p p e a r ed a f t er s p r a y i n g. E x t r a c ts of f r e sh t u b er t h at h a s n 't b e en i n o c u l a t ed h a ve s o me of t h e se m e t a b o l i t e s. If we let t he t i s s ue a g e, i n d e ed we s ee t h at a f ew n ew m e t a b o l i t es a p p e ar and o t h e rs i n c r e a s e. If we i n o c u l a te R u s s e tt B u r b a nk w i th P. i n f e s t a ns ( s u s c e p t i b l e ), we s ee v e ry l i t t le i n c r e a se in t he t e r p e n o i d - t y pe or s t e r o i d - t y pe c o m p o u n ds as c o m p a r ed to t he a g ed u n i n o c u l a t ed t u b e r s. r e- s i s t a nt v a r i e t y, we s ee a v e ry m a r k ed b u i l d - up of m e t a b o l i t e s. I t 's t e m p t i ng to s ay t h en t h at K e n n e b ec h as g e n e t ic i n f o r m a t i on w h i ch is c o n t r o l l i ng t he s y n t h e s is of t h e se c o m p o u n ds and t h at t he R u s s et B u r b a nk d o es not. A c t u a l ly t h a t 's a m i s l e a d i ng s t a t e m e n t. t he n o n - p a t h o g en of p o t a t o, H. c a r b o n u m, we f i nd t h at both c u l t i v a rs a re e q u a l ly a b le to s y n t h e s i ze t h e se t e r p e n o i ds v e ry r a p i d ly a f t er i n f e c t i o n s. We c an not s t a te t h at R u s s et B u r b a nk d o e s n 't h a ve t he g e n e t ic i n f o r m a t i on to p r o d u ce t h e se c o m p o u n d s. A ll we c an s ay is t h at in t he p r e s e n ce of a r a ce w h i ch c an a t t a ck If we i n o c u l a te R u s s et B u r b a nk and K e n n e b ec w i th If we i n o c u l a te t he K e n n e b e c, t he t he R u s s e tt B u r b a nk ( f or s o me r e a s on t he g e n e t ic i n f o r m a t i on is not being e x p r e s s e d) w h i ch would l e ad to t he p r o d u c t i on of t h e se c o m p o u n d s. I t 's a l so p o s s i b le t h at t he c o m p o u n ds a re being d e- g r a d ed in t he c o m p a t i b le i n t e r a c t i o n. What is t he r e l a t i o n s h ip b e t w e en t he t e r p e n o i ds and r e s i s- t a n c e? We f i nd t h at t h e re a re a n u m b er of t e r p e n o id c o m p o u n ds t h at c o n s i s t e n t ly a r i se in i n c o m p a t i b le or r e s i s t a nt r e a c t i o ns w h i ch we do not f i nd in t he s u s c e p t i b le or c o m p a t i b le r e a c t i o n s. T h e se c o m p o u n ds a re i n h i b i t o ry to P. i n f e s t a n s. T h ey a re a l so to H. c a r b o n u m, and t h ey a c c u m u l a te v e ry r a p i d ly a f t er i n f e c t i on to l e v e ls w h i ch c an a c c o u nt f or t he c o n t a i n m e nt of t he o r g a n i sm in t he p l a n t. d i s e a se r e s i s t a n ce m e c h a n i sm in a n i m a l s. We d o n 't h a ve c o m- p l ex p r o t e in a n t i b o d i es p r o d u c ed or i n t e r f e r o n - t y pe s u b s t a n c e s, but we do h a ve s i m p l er c o m p o u n d s. T h ey m ay be a n d / or p h e n o l s, d e p e n d i ng on t he plant i n f e c t e d. T h e se c o m p o u n d s, in t he c a se of p l a n t s, m ay h a ve a v e ry i m p o r t a nt r o le in d i s e a se r e s i s t a n c e. l e v e ls w h i ch inhibit f u r t h er d e v e l o p m e nt of t he i n f e c t i o us a g e n t. It a p p e a rs we h a ve a m e c h a n i sm w h i ch r e s e m b l es a In r e s i s t a nt t i s s u e s, t h ey build up v e ry q u i c k ly to i n h i b i t o ry t e r p e n o i ds M e c h a n i s ms t h at A c t i v a te and S u p p r e ss D i s e a se R e s i s t a nt R e a c t i o ns T h e se c o m p o u n d s, w h i ch build up in r e s i s t a nt t i s s u e s, a re sometimes called phytoalexins, One th.at we've worked with, in p o t a to is t he c o m p o u nd r i s h i t i n. We h a ve a l so i s o l a t ed a n o t h er c o m p o u n d, p h y t u b e r i n. R i s h i t in b e c o m es evident v e ry soon ( 24 h o u r s) a f t er i n o c u l a t i on and r a p i d ly a c c u m u l a t e s. P h y t u- b e r in a l so b u i l ds up in t he i n c o m p a t i b le or r e s i s t a nt r e a c t i o n. H ow r e l i a b le is t h is a c c u m u l a t i o n? We h a ve u s ed e l e v en c u l- t e s t ed t i v a rs of p o t a t os and s ix r a c es of t he f u n g u s. We h a ve n u m b e r o us c o m p a t i b le and i n c o m p a t i b le i n t e r a c t i o n s. In a ll t h e se c a s e s, r i s h i t in and p h y t u b e r in r a p i d ly a c c u m u l a te in t he i n c o m p a t i b le but not t he c o m p a t i b le or s u s c e p t i b le i n t e r a c t i o n. We f e el c o n f i d e nt i n d e ed t h at t h is is a r e al thing and t h e se t e r p e n o i ds a re a s s o c i a t ed w i th t he d i s e a se r e s i s t a n ce r e a c t i o n. two What is it t h at is i n d u c i ng t he s y n t h e s is and t he a c c u m u l a- Is it s o m e t h i ng i n d u ce l i v- t i on of r i s h i t in in t he i n c o m p a t i b le i n t e r a c t i o n? v e ry s p e c i f ic t h at a r i s es only in t he i n t e r a c t i o n? C an we t he p l a nt to p r o d u ce and a c c u m u l a te t he r i s h i t in without t he ing f u n g u s? We f o u nd t h at a c e l l - f r ee s o n i c a te of a ny r a ce of t he f u n g us c an i n d u ce t he p r o d u c t i on of r i s h i t in and p h y t u b e r in in t he i n c o m- p o t a to t u b er to l e v e ls e v en g r e a t er t h an t h o se f o u nd in an p a t i b le i n t e r a c t i o n. I n t e r e s t i n g ly enough, e v en c u l t i v a rs " l a c k- ing g e n es f or r e s i s t a n c e ," a c c u m u l a te r i s h i t in and p h y t u b e r in w h en t r e a t ed w i th s o n i c a t es of P. i n f e s t a n s. R u s s et B u r b a nk is a c u l- t i v ar t h at is s u s c e p t i b le to a ll t he k n o wn r a c es of P. t h e r e f o r e, it would be t e m p t i ng to s ay t h at it d o es not h a ve m a j or R g e n es f or r e s i s t a n ce to P. i n f e s t a n s. H o w e v e r, a s o n i c a te f r om any r a ce of P. i n f e s t a ns i n d u c es r a p id a c c u m u l a t i on of b o th t e r p e n o i d s. T he a c c u m u l a t i on would not h a ve o c c u r r ed if t he p o t a to had b e en i n o c u l a t ed with a c o m p a t i b le r a c e. How s p e c i f ic is t h is r e s p o n se of t he p o t a to to P. i n f e s t a ns ? We c an i n d u ce t he a c c u m u l a t i on of r i s h i t in and p h y t u b e r in with a b r o ad s p e c t r um of f u n gi i n c l u d i ng H. c a r b o n u m. W e ' re not about a s p e c i f ic a n t i b o d y - a n t i g e n - t y pe of r e l a t i o n s h i p. W e ' re t a l k i ng about a b r o ad s p e c t r um of c o m p o u n ds w h i ch a re r a t h er n o n - s p e c i f ic but w h i ch t he f u n g us w i ll e i t h er i n d u ce t he a c- c u m u l a t i on of or not, d e p e n d i ng u p on w h e t h er it c an or c a n n ot a t t a ck t h at p a r t i c u l ar h o s t. One w o n d e rs t h en what is it t h a t 's If t he h a p p e n i ng in t he c o m p a t i b le or s u s c e p t i b le i n t e r a c t i o n. p o t a to t h at is s u s c e p t i b le h as t he p o t e n t i al f or p r o d u c i ng t he a n t i b i o t ic c o m p o u n d s, why is it t h at t h is p o t e n t i al is not e x p r e s s e d; t h at t h is p o t e n t i al is not r e a l i z e d? T h e re a re s e v e r al p o s s i b i l i- t i e s. One, is t h at in t he s u s c e p t i b le i n t e r a c t i o n, t h e se a n t i b i o t ic c o m p o u n ds a re p r o d u c ed but t h e y ' re r a p i d ly d e t o x i c a t e d. T h ey a re b r o k en down into n o n - f u n g i t o x ic c o m p o u n d s. A n o t h er p o s s i- b i l i ty is t h a t, f or s o me r e a s on in t he s u s c e p t i b le t he f u n g us is s u p r e s s i ng t he s y n t h e s is or a c c u m u l a t i on of t h e se c o m p o u n d s. r e s i s- t a n ce m e c h a n i s m, t h at it is a b le to a t t a ck t he p o t a t o. It is b e c a u se of t he s u p r e s s i on of a g e n e r al i n t e r a c t i o n, i n f e s t a n s; t he t a l k i ng We s et up a s e r i es of e x p e r i m e n ts i n o c u- l a t ed with t he c o m p a t i b le r a c e, w a i t ed a p e r i od of 12 or 24 h o u r s, and t h en i n o c u l a t ed with t he i n c o m p a t i b le r a c e. I n- o c u l a t i on with t he i n c o m p a t i b le r a ce n o r m a l ly l e a ds to t he a c c u m u l a t i on of r i s h i t in and p h y t u b e r i n. What we f o u nd w as in w h i ch we t h at w h en we p r e c e d ed t he i n c o m p a t i b le r a ce w i th t he c o m p a t i b le r a ce by 12 or 24 h o u r s, we s u p p r e s s ed t he r e s p o n se to t he i n c o m- p a t i b le r a c e. S u s c e p t i b i l i t y, at l e a st in t h is s y s t e m, w as b a s ed on s o m e h ow s u p p r e s s i ng a r e s i s t a n ce r e s p o n s e. Not a l a ck of a p o t e n t i al f or p r o d u c i ng t he c o m p o u n d, not a l a ck of p o t e n t i al f or i n d u c i ng i ts p r o d u c t i o n, but r a t h er an a c t i ve m e c h a n i sm f or s u p p r e s s i o n. If we h a ve i n o c u l a t i on w i th an i n c o m p a t i b le f o l l o w ed by a c o m- p a t i b le r a c e, t he i n c o m p a t i b le r a ce c o m p l e t e ly p r o t e c ts t he p o t a to a g a i n st s u b s e q u e nt i n o c u l a t i on by t he c o m p a t i b le r a c e. it f i r st w i th t he r e s i s t a nt c o m b i n a t i o n, we get t he build up of r i s h i t in and s t a rt a s e r i es of m e t a b o l ic e v e n ts w h i ch p r o t e ct p o t a t o. w a it a p e r i od of 12 h o u rs and i n o c u l a te it w i th t he r a c e, r i s h i t in a c c u m u l a t i on is m a r k e d ly r e d u c ed and we a l so d o n 't h a ve p r o t e c t i o n. We h a v e, in f a c t, s u s c e p t i b i l i t y. A 2 4 - h o ur lag b e t w e en i n o c u l a t i on w i th t he c o m p a t i b le and i n c o m p a t i b le r a ce i n c r e a s es If we i n o c u l a te t he p o t a to w i th t he c o m p a t i b le r a c e, i n c o m p a t i b le If we hit t he t h en s u p p r e s s i o n. Now what h a p p e ns on t he m e t a b o l ic l e v el w h en we do t h i s? M e t a b o l i c a l l y, how do we e x p l a in t h i s? We f i nd an i n t e r a c t i o n^ t h en t he a c c u m u l a t i on of r i s h i t in is s u p r e s s ed by a c o m p a t i b l e. A n o t h er t e r p e n o id a c c u m u l a t es w h i ch is not f u n g i t o x i c. It a p p e a rs t h at by s u p p r e s s i ng r i s h i t in a c c u m u l a t i o n, w e ' ve c a u s ed t he a c c u- m u l a t i on of a n o t h er c o m p o u nd in t he p a t h w a y. T h is c o m p o u nd is not t o x ic to t he f u n g u s. T he f u n g us c an g r o w. I n o c u l a t i on w i th a c o m p a t i b le r a ce 12 to 24 h o u rs b e f o re t r e a t m e nt w i th a f u n g al s o n i c a te a l so s u p p r e s s es a c c u m u l a t i on of r i s h i t in and p h y t u b e r i n. I t 's not a m a t t er of one r a ce c o m p e- t i ng w i th a n o t h e r, but is in f a ct t h at t he c o m p a t i b le or s u s c e p t i b le c o m b i n a t i on i n t e r f e r es with t he r e s p o n se to t he r a ce or i ts p r o d u c t s. i n c o m p a t i b le Not only h as t he a c c u m u l a t i on of r i s h i t in and p h y t u b e r in b e en s u p p r e s s e d, but t he n e c r o s is a s s o c i a t ed h y p e r s e n s i t i ve r e a c t i on or r e s i s t a nt r e a c t i on is a l so not e v i d e n t. F or s o me r e a s o n, now t he p o t a to m ay r e c o g n i ze t he i n c o m p a t i b le r a ce as c o m p a t i b l e. T h is is a bit of t he p o t a to s t o r y. And i n d e ed we s ee t h at c o m p o u n ds a re p r o d u c ed in r e s i s t a nt r e a c t i o n s. T h e se c o m p o u n ds a re i n h i b i t o r y. T h ey a re p r o d u c ed at l e v e ls t h at c an inhibit m i c r o o r- g a n i s m s. We c an i n d u ce t h e ir p r o d u c t i on w i th c h e m i c a ls p r o- d u c ed by t he f u n g u s. a p p e ar d ue to t he a b i l i ty or i n a b i l i ty to p r o d u ce a c o m p o u nd by t he p l a n t, but r a t h er w h e t h er it h a p p e ns in an i n t e r a c t i on s o on e n o u gh and in s u f f i c i e nt m a g n i t u d e. In t h is d i s e a s e, s u s c e p t i b i l i ty d o es not D i s e a se S u s c e p t i b le a nd R e s i s t a nt I n t e r a c t i o ns in G r e en B e a ns l i n d e m u t h i a n u m. Is it r e a l ly a c c u r a t e, h o w e v e r, to s ay t h at L e t 's t a ke a look at a n o t h er i n t e r a c t i o n. T h is is t he In t he c a se i t!s an a e s t h e t ic l o s s. T h is is not, h o w e v e r, i n t e r- a c t i on of g r e en b e an w i th C o l l e t o t r i c h um I n o- c u l a te s e e d l i n gs of t he s u s c e p t i b le c u l t i v ar R ed K i d n ey w i th t he b e ta and g a m ma r a ce of t he C. l i n d e m u t h i a n u m, and a s u s c e p t i b le r e a c t i on is a p p a r e n t. t h is is a s u s c e p t i b le p l a n t? We s ay it is s u s c e p t i b le b e c a u se is s u f f i c i e nt d a m a ge to c a u se s o me e c o n o m ic l o s s. of an o r n a m e n t a l, a b i o c h e m i c al d e f i n i t i o n. W h en we look at t he p l a n t, we s ee not an a c c u r a te d e f i n i t i on by a ny m e a n s. E v en in t he s u s c e p t i b le i n t e r a c t i o n, we h a ve a r e as of v e ry r e al and v e ry m a r k ed r e s i s- t a n c e. As we a p p r o a ch t he r o ot z o n e, t he l e s i o ns b e c o me v e ry r e s t r i c t e d. D o wn in t he r o ot a r ea i t s e lf we f i nd t y p i c al h y p e r- s e n s i t i v e - t y pe r e a c t i o n s. T he o r g a n i sm g e r m i n a t es in t h e se a r e a s, p e n e t r a t e s, but you get a h y p e r s e n s i t i ve r e a c t i on - -a v e ry r e s i s t a nt r e a c t i o n. E v en t h is s u s c e p t i b le p l a n t, s u s c e p t i b le to t he b e ta and g a m ma r a c es of f u n g u s, h as t i s s u es t h at a re e x p r e s s i ng h y p e r s e n s i t i ve t h e re i tfs r a p id r e s i s t a n c e. If we a s s u me t he g e n e t ic i n f o r m a t i on is t he s a me in a ll t he c e l ls of t he p l a n t, i t 's t e m p t i ng to s ay t h at t h is p l a n t, i n d e e d, h as g e n e t ic i n f o r m a t i on f or r e s i s t a n ce and in one p a rt i t 's b e i ng e x p r e s s ed and a n o t h er i t 's not. I t 's not a m e c h a n i c al m a t t er b e c a u se t he o r g a n i sm g e r m i n a t es and p e n e t r a t e s. v e ry n i ce to get to t he k ey f or t he e x p r e s s i on t h a t 's so e f f e c t i ve in r o o ts and s o m e h ow u se t h is k ey to e x p r e ss t he g e n es in a ll t i s s u e. It would be In t he b e an - C. l i n d e m u t h i a n um i n t e r a c t i o n, as w i th p o t a t o- P. i n f e s t a n s, we h a ve c u l t i v a rs r e s i s t a nt to s o me r a c es and s u s- c e p t i b le to o t h e r s. We c an t a ke a c u l t i v a r, t he g e n e t ic i n f o r m a t i on of t he p l a nt being c o n s t a n t, a nd v a ry t he r a ce of t he f u n g u s. Or we c an t a ke s e v e r al c u l t i v a rs w h i ch r e a ct d i f f e r e n t ly to a s i n g le r a c e. T h is is a b e a u t i f ul s y s t em to c r o s s c h e ck a ll of o ur w o rk a nd a ll t he w o rk I d e s c r i be w i th s e e d l i n gs is v a l id f or p l a n ts and p o d s. T he s a me c u l t i v ar m ay c h a n ge f r om c o m p l e te r e s i s t a n ce to c o m p l e te s u s c e p t i b i l i ty d e p e n d i ng on t he r a ce of f u n g us u s ed f or i n o c u l a t i o n. T h is a r g u es v e ry s t r o n g ly a g a i n st t he t h o u g ht t h at a r e s i s t a nt p l a nt h as g e n es f or doing s o m e t h i ng and a s u s- c e p t i b le p l a nt d o es not. H e re we s ee t he s a me p l a nt r e a c t i ng one w ay to one r a ce and a n o t h er w ay to a n o t h er r a c e. in a l i t t le m o re d e t a i l. L e t 's look at t h e se i n t e r a c t i o ns If we i n o c u l a te a s u s c e p t i b le or r e s i s t a nt c u l t i v ar w i th t he s p o r es of C. l i n d e m u t h i a n u m, t he s p o r es g e r m i n a te w i t h in 3 to 5 h o u rs a nd g e r m i n a te e q u a l ly w e ll on a s u s c e p t i b le or r e s i s t a nt c u l t i v a r. About f i ve h o u rs a f t er g e r m i n a t i o n, a t h i ck w a l l ed a p p r e s o r i um f o r m s. T h is is a s t r u c t u re f r om w h i ch t he o r g a n i sm w i ll e m e r ge a g a in and t h en p e n e t r a te i n to t he p l a n t. T h is d o e s n 't h a p p en i m- m e d i a t e l y. About 36 to 48 h o u rs a f t er i n o c u l a t i on t he o r g a n i sm e m e r g es a g a in f r om t he a p p r e s o r i um and p e n e t r a t es into t he p l a n t. Up u n t il t h is t i m e, t he r e a c t i on is i d e n t i c al on t he s u s c e p t i b le a nd t he r e s i s t a nt p l a n t. T h i r t y - s ix to f o r t y - e i g ht h o u rs a f t er i n o c u l a t i o n, w h en t he f u n g us h as p e n e t r a t ed into t he r e s i s t a nt p l a n t, we s ee a g r a n u l a t i on of t he c y t o p l a sm -- a h y p e r s e n s i t i ve r e a c t i on -- and t he f u n g us is c o n t a i n e d. T he f u n g us g o es no f u r t h er t h an one or t wo c e l l s. At t h at s a me t i m e, in t he s u s c e p- t i b le i n t e r a c t i o n, t he o r g a n i sm h as g r o wn c o n s i d e r a b ly m o re but t h e re h as not b e en a g r a n u l a t i on of t he c y t o p l a s m, t he c e l ls a re i n t a c t. T h is is t r u e, e v en t h o u gh e r a b ly m o r e. A f t er 9 6 h o u r s, t he f u n g us h as r a m i f i ed t he t i s s ue and s o me c e ll w a l ls a p p e ar d i s o r g a n i z e d. N i n e t y- s ix h o u r s, or p e r h a ps 110 h o u rs a f t er i n o c u l a t i o n, in t he s u s c e p t i b le i n t e r a c t i o n. R e s i s t a n c e, t h e r e f o r e, a v e ry h i gh d e g r ee of i n c o m p a t i b i l i t y, w h e r e as is a h i gh d e g r ee of c o m p a t i b i l i t y. t he f u n g us and t he p l a nt get a l o ng f or a long p e r i od of t i m e, u n t il t he o r g a n i sm h as w o rn out i ts w e l c o m e. Only t h en d o es i n j u ry o c c u r. W h a t 's h a p p e n i ng c h e m i c a l ly in t h is t i s s u e? T o t al p h e n o ls a c c u m u l a te in b e an s e e d l i n gs a f t er i n o c u l a t i on w i th a l e s i o ns a p p e ar r e p r e s e n ts s u s c e p t i b i l i ty In t he s u s c e p t i b le i n t e r a c t i on t he f u n g us h as g r o wn c o n s i d- t h r o u gh c o m p a t i b le or i n c o m p a t i b le r a ce of C. l i n d e m u t h i a n um or a n on p a t h o g en of b e a n, H. c a r b o n u m. In t he i n c o m p a t i b le r e a c t i on of t he g a m ma r a ce of C. l i n d e m u t h i a n u m, h o w e v e r, t he build up of p h e n o l ic c o m p o u n ds s t a r ts about 48 h o u rs a f t er i n o c u l a t i on and p r o c e e ds t h r o u g h o ut t he t i me i n t e r v al we u s ed f or o ur study. In t he s u s c e p t i b le or c o m p a t i b le i n t e r a c t i o n, we d o n 't get t h is build up e a r l y, but only w h en l e s i o ns t y p i c al of t he s u s c e p t i b le i n t e r a c t i on s t a rt to a p p e a r. T he s u s c e p t i b le and r e s i s t a nt i n- t e r a c t i on a p p e ar to be doing t he s a me t h i n g. T he r e s i s t a nt i n t e r- a c t i on d o es it v e ry q u i c k l y; t he s u s c e p t i b le d o es it m u ch m o re slowly. T he t o t al p h e n ol a c c u m u l a t i on in both r e a c t i o ns is about t he s a m e. P e r h a ps an a n a l o gy h e re would be a c a se of s m a l l p o x. A t r e m e n- p e r s on who h as n e v er b e en i m m u n i z ed a g a i n st s m a l l p ox c o n t a c ts s m a l l p ox and is dying, but is p r o d u c i n g, j u st b e f o re d e a t h, dous q u a n t i t i es of a n t i b o d i e s. U n f o r t u n a t e l y, t he r e s p o n se m ay be too l a t e. T he p e r s on who h as b e en i m m u n i z ed c an r e a ct i m- m e d i a t e ly to s m a l l p ox and t he i n f e c t i o us a g e nt n e v er g e ts to do If we i n o c u l a te w i th H. c a r b o n u m, a v e ry m a r k ed any d a m a g e. i n c r e a se in p h e n o ls o c c u rs in 12 to 24 h o u r s, and t h is h i gh l e v el t a l k- is m a i n t a i n ed t h r o u g h o ut t he t e s t. T he c o m p o u nd t h at w e ' re ing about p a r t i c u l a r ly in t h is i n t e r a c t i on is t he c o m p o u nd p h a s e- ollin. W e ' re not l i m i t ed to t h is c o m p o u nd and we o b s e r ve a s e r i es of c o m p o u n ds t h at c o n t r i b u te to t he r e s i s t a n ce m e c h a n i s m. P h a s e o l l i n, one of t he a n t i b i o t ic c o m p o u n ds p r o d u c ed by t he g r e en b e an in r e s p o n se to i n f e c t i o n, and r i s h i t i n, w h i ch is p r o d u c ed t he p o t a to i n t e r a c t i o n, a re v e ry d i f f e r e nt c o m p o u n d s. T he p a t t e rn of i n c r e a s ed a c c u m u l a t i on of p h e n o ls m ay a l so hold t r ue f or e n z y m es c o n t r o l l i ng t he s y n t h e s is of t h e se c o m p o u n d s. in A p o s s i b le k ey e n z y me in t he s y n t h e s is of p h a s e o l l i ns in t he r e s i s t a nt l y a s e. A v e ry m a r k ed i n c r e a se in t he p h e n y l a l a n i ne is a m m o n ia a c t i v i ty of t h is e n z y me o c c u rs 48 h o u rs a f t er i n o c u l a t i o n. An i n c r e a se a l so o c c u rs in t he s u s c e p- t i b le i n t e r a c t i o n, but it c o m es m u ch l a t e r. T h is f o l l o ws t he p a t t e rn of p h e n o l ic s y n t h e s is and a c c u m u l a t i o n. T h e r e f o r e, an e n z y m e, . t he c a t a l y st f or t he s y n t h e s is of t h e se c o m p o u n d s, v e ry c l o s e ly f o l l o ws t he p a t t e rn f or a c c u m u l a t i on of t he c o m p o u n ds t h e m s e l v e s. A v e ry m a r k ed i n c r e a se in t he e n z y m e 's a c t i v i ty is e v i d e nt 12 h o u rs i n t e r a c t i on 36 to a f t er b e an s e e d l i n gs w e re i n o c u l a t ed w i th H. c a r b o n u m. Activity- r e t u r ns to t he l e v el in u n i n o c u l a t ed t i s s ue 12 to 24 h o u rs S e v e r al c o m p o u n d s, o t h er t h an p h a s e o l l i n, a c c u m u l a te in t i s s ue i n o c u l a t ed with H. c a r b o n u m. l a t e r. E s t a b l i s h m e nt of D i s e a se R e s i s t a nt S t r a i ns It a p p e a rs a s i n g le c o m p o u nd is not r e s p o n s i b le f or d i s e a se r e s i s t a n c e, but r a t h er a s p e c t r um of c o m p o u n ds is i n v o l v e d. T h ey m ay be r e l a t e d, but t h ey a re d i f f e r e n t. T h is h e l ps e x p l a in why in n a t u re r e s i s t a n ce is t he r u le arid s u s c e p t i b i l i ty is Ahe e x c e p t i o n. When we u se a f u n g i c i de to c o n t r ol d i s e a s e, we v e ry o f t en p i ck up r e s i s t a nt s t r a i ns b e c a u se w e ' re d e a l i ng w i th a s i n g le c o m- pound. T he c h a n c es of m u t a t i ng a r o u nd a s i n g le c o m p o u nd a re i n f i n i t e ly g r e a t er t h an t he c h a n c es of m u t a t i ng a r o u nd a s e r i es of c o m p o u n ds t h at a re p r o d u c ed in a n a t u r al r e s i s t a n ce m e c h- a n i s m. If t he t h i n gs I ' ve s a id a b o ut d i s e a se r e s i s t a n ce a re t r u e, and of c o u r se I b e l i e ve t h ey a r e, t h en we s h o u ld be a b le to i m m u n i ze p l a n t s. We s h o u ld be a b le to p r o t e ct a b e an as we did t he p o t a to by i n o c u l a t i ng with a r a ce of p a t h o g en w h i ch c a n n ot a t t a ck t h at c u l t i v ar and p r o t e ct it a g a i n st a r a ce t h at can. We s h o u ld a l so be a b le to u se a n on p a t h o g en -- i n o c u l a te a p l a nt w i th a non p a t h o g en to p r o t e ct it a g a i n st a p a t h o g e n. T he b e an c u l t i v a r, P e r ry M a r r o w, is s u s c e p t i b le to t he g a m ma and r e s i s t a nt to t he b e ta r a ce of C. l i n d e m u t h i a n u m. i n o c u l a t ed with t he g a m ma r a ce s h ow l a r ge c o a l e s c ed S e e d l i n gs f i r st i n o c u l a t ed w i th t he b e ta r a ce and 24 h o u rs i n n o c u l a t ed w i th t he g a m ma r a ce a re a b s o l u t e ly p r o t e c t e d. T h e r e f o r e, by i n o c u l a t i ng w i th t he r a ce t h at cannot a t t a ck b e- f o re we i n o c u l a te w i th t he r a ce t h at can, we c an p r o t e ct t he b e a n. A v e ry h i gh d e g r ee of p r o t e c t i on a g a i n st a r a ce of t he p a t h o g en t h at c an a t t a ck is i n d u c ed by p r i or i n o c u l a t i on w i th H. c a r b o n um or an A l t e r n a r ia s p e c i e s, both non p a t h o g e ns of b e a n. S e e d l i n gs l e s i o n s. l a t er We c an do t he s a me t h i ng w i th a r e c i p r o c al r a ce d i f f e r e n t i al u s i ng a c u l t i v ar s u s c e p t i b le to t he b e ta and r e s i s t a nt to t he g a m ma r a c e. A f t er i n o c u l a t i on w i th t he b e ta r a c e, we s ee l e s i o n s. If we i n o c u l a te with t he g a m ma r a c e, wait 24 h o u r s, and i n o c u l a te w i th t he b e t a, we get c o m p l e te p r o t e c t i o n. We a l so s ee a v e ry h i gh d e g r ee of p r o t e c t i on u s i ng t he two non p a t h o g e ns of b e a n. T h is c an be r e p e a t ed u s i ng leaf t i s s ue and p o d s. t y p i c al A n i m a ls a re i m m u n i z ed a g a i n st i n f e c t i o us a g e n ts by u s i ng If t he m e c h a n i s ms f or p r o- t he a t t e n u a t ed a g e n t; i . e ., t he i n f e c t i o us a g e nt is t r e a t ed so t h at it is u n a b le to f u n c t i on in c a u s i ng d i s e a se but s t i ll is a b le to t he i m m u ne r e s p o n se in t he a n i m a l. t e c t i on in p l a n ts and a n i m a ls a re s i m i l a r, we s h o u ld be a b le to do t he s a me t h i ng in o ur p l a nt s y s t e m. W e ' ve a c c o m p l i s h ed t h is by t a k i ng a d v a n t a ge of t he f a ct t h at C. l i n d e m u t h i a n um is s e n s i t i ve t e m p e r a t u r es f r om 37°C. to 40°C. At t h e se t e m p e r a t u r es p l a nt is u n h a r m ed but g r o w th of C. l i n d e m u t h i a n um i n o c u l a t ed b e an p l a n ts w i th a r a ce of C. l i n d e m u t h i a n um a t t a ck and let t he f u n g us d e v e l o p. T he p l a n ts w e re t h en h e a t- t r e a t ed and r e - i n o c u l a t ed with t he s a me r a c e. T h e re w e re no s y m p t o ms and t he f u n g us w as c o n t a i n e d. a nd t h en i n o c u l a t e, we a c t u a l ly i n c r e a se s u s c e p t i b i l i t y. T he p r o t e c t i o n, t h e r e f o r e, is not due to t he h e at t r e a t m e n t, p er s e. I t 's due to t he h e at t r e a t m e nt of t he f u n g us in t he p l a n t. If we h e at t r e at a l o n e, i n d u ce to t he b e an is s t o p p e d. We t h at c an How f ar f r om t he s i te of i n d u c t i on c an we d e m o n s t r a te p r o- t e c t i o n? P l a n ts d i s c u s s ed e a r l i er w e re p r o t e c t ed by b e i ng s p r a y ed with i n o c u l u m. To d e m o n s t r a te d i s t a n ce e f f e c t s, d r o ps of i n d u c er w e re p l a c ed on h y p o c o t y ls at s o me d i s t a n ce f r om t he c h a l l e n ge i n o c u l u m. In s o me c a s e s, i n d u c er and c h a l l e n ge w e re a p p l i ed at t he s a me t i m e; in o t h er c a s e s, i n d u c er p r e c e e d ed c h a l l e n ge by 12 to 24 h o u r s. W h en a t i me i n t e r v al of 12 h o u rs s e p a r a t ed i n d u c er and I t 's not t h at n o- c h a l l e n g e, p l a n ts w e re c o m p l e t e ly p r o t e c t e d. t h i ng h a p p e n e d, but r a t h er t h at we o b s e r v ed a t y p i c al h y p e r- s e n s i t i ve r e a c t i o n. A m e s s a ge m u st h a ve gone f r om t he i n d u c er s i te to t he c h a l l e n ge s i te - -a d i s t a n ce of 0.5 to 1.0 c m. T he m e s- s a ge s i g n a l ed t he p l a nt to r e c o g n i ze t he c o m p a t i b le r a ce as c o m p a t i b l e. i n- In t he n e xt s e r i es of e x p e r i m e n t s, we put s p o r es of an i n c o m- .22 m i c r on m i l l i p o re f i l t e r. T he r e s u l t i ng d i f f u s a te w as s t e r i l e. p a t i b le r a ce on h y p o c o t y ls as l i t t le d r o p s. T he d r o ps w e re c o l- l e c t ed a f t er a b o ut 48 h o u rs and f u n g us r e m o v ed by f i l t r a t i on t h r o u gh a T he d i f f u s a te w as p l a c ed on a n o t h er h y p o c o t yl and t he s i te l a t ed w i th a r a ce of t he f u n g us t h at s h o u ld c a u se d i s e a s e. T he p l a n ts t r e a t ed t h is w ay w e re c o m p l e t e ly p r o t e c t e d. D i f f u s a t es f r om c o m p a t i b le i n t e r a c t i o ns did not p r o t e c t. T he living i sm is no l o n g er n e e d ed to i n d u ce r e s i s t a n ce and we c an p r o t e ct w i th a c h e m i c al f a c t or p r o d u c ed in t he i n c o m p a t i b le i n t e r a c t i o n. T he c h e m i c al i n d u c i ng r e s i s t a n ce is not i t s e lf i n h i b i t o ry to t he f u n g u s. o r g a n- i n o c u- What h a p p e ns if we i n o c u l a te w i th a non p a t h o g en of b e a n? C an we do t he s a me t h i n g? T he a n s w er i s, y e s. We c an p r o t e ct with m a ny C o l l e t o t r i c h um s p e c i es n on p a t h o g e n ic on b e a n. T he p i c t u re is not as s i m p le as I ' ve e x p l a i n ed it but it b e c o m es e v en m o re f a s c i n a t i ng w h en we look at j u st one o t h er p i e ce of i n f o r m a t i o n. P r o t e c t i on w i th t he d i f f u s a t es d e p e n ds not only u p on t he p r e s e n ce of i n d u c er in t he d i f f u s a t e, but a l so u p on t he p r e s e n ce of g e n e t ic r e c e p t or c i t es in t he p l a n t. C u l t i v a rs a re p r o t e c t ed by d i f f u s a t es f r om i n c o m p a t i b le t he t i o ns only if t h ey a re r e s i s t a nt to t he r a ce u s ed to p r e p a re d i f f u s a t e. r e a c- T h e r e f o r e, d i f f u s a te f r om t he i n c o m p a t i b le r e a c t i on of t he b e ta P e r ry M a r r ow and g a m ma r a ce on t he c u l t i v a r, P e r ry M a r r o w, r e s i s t a nt to t he b e ta r a c e, w i ll p r o t e ct P e r ry M a r r ow a g a i n st t he g a m ma r a c e. T he d i f f u- s a te w i ll not p r o t e ct t he c u l t i v ar T op C r o p, s u s c e p t i b le to t he b e ta r a c e, a g a i n st t he b e ta r a c e. T he r e c i p r o c al s i t u a t i on is e q u a l ly t r u e. M i x ed d i f f u s a te f r om t he b e ta r a ce on r a ce on T op C r op w i ll p r o t e ct both c u l t i v a rs a g a i n st t he g a m ma and b e ta r a c e s, r e s p e c t i v e l y. T e n n e s s ee G r e en P od is r e s i s t a nt to t he b e ta a nd g a m ma r a c e s, but it is s u s c e p t i b le to t he d e l ta r a ce of C l i n d e m u t h i a n u m. D i f f u s a t es f r om T e n n e s s ee G r e en P od i n o c u l a t ed w i th t he b e ta or g a m ma r a ce p r o t e ct a g a i n st t he d e l ta r a c e. D i f f u s a te f r om t he b e ta r a ce on T e n n e s s ee G r e en P od w i ll a l so p r o t e ct c u l t i v a r s, s u ch as P e r ry M a r r o w, r e s i s t a nt to t he b e ta r a c e, a g a i n st t he g a m ma r a c e. T he s a me d i f f u s a te w i ll not p r o t e ct c u l t i v a r s, s u ch as T op C r o p, a g a i n st t he b e ta r a c e. T he r e c i p r o c al s i t u a t i on is e q u a l ly t r u e. T h is is v e ry good e v i d e n ce t h at not only a f a c t or f or i n d u c- t i on is n e c e s s a ry f or p r o t e c t i o n, but a g e n e t i c a l ly c o n t r o l l ed r e- c e p t or s i te m u st be a v a i l a b le to get t he m e s s a g e. What h a p p e ns if we i n o c u l a te w i th a non p a t h o g en of b e an l i ke Co 11 e t o t r i c h um t r u n c a t u m? A ll g r e en b e an c u l t i v a rs h a ve t he r e c e p t or s i te f or r e s i s t a n ce to C. t r u n c a t um and d i f f u s a te f r om C. t r u n c a t um on g r e en b e an w i ll p r o t e ct a ny g r e en b e an c u l t i v ar a g a i n st t he b e t a, g a m ma or d e l ta r a ce of C. e v en p r o t e c ts g r e en b e an c u l t i v a rs t h at a re s u s c e p t i b le to a ll t he k n o wn r a c es of C. l i n d e m u t h i a n u m. T he c u l t i v a rs s u s- c e p t i b le to a ll t he k n o wn r a c es of C. l i n d e m u t h i a n um h a ve r e c e p t or s i te f or r e s i s t a n ce to C. l i n d e m u t h i a n u m. t r u n c a t u m. In f a c t, C. t he t r u n c a t um t he t h at T h is of c o u r se is a ll v e ry e x c i t i ng to us b e c a u se it f o l l o ws r e s i s- I n d e e d, r e s i s t a n ce is g e n e t i c a l ly c o n t r o l l e d, as a re a ll a p a t t e rn w h i ch f o l l o ws t he g e n e t i cs of s u s c e p t i b i l i ty and t a n c e. O n ce a g a in I e m p h a s i ze t he i m p o r t a n ce of e x p r e s s i ng g e n e t ic i n f o r m a t i on f or r e s i s t a n c e. T h is d o e s n 't m e an t h at g e n e t i c i s ts w e re w r o ng in t h e ir t h o u g h ts c o n c e r n i ng r e s i s- t a n c e. t h i n g s. But t he t h i n k i ng as we s ee it is t h at i t 's t he r e g u l a t o ry g e n es t h at a re c o n t r o l l i ng t he e x p r e s s i on of t he s t r u c t u r al g e n es b e c o me i m p o r t a nt in m a ny c a s es of r e s i s t a n c e. t he q u e s t i on is not w h e t h er a p l a nt c an do s o m e t h i ng but w i ll it do s o m e t h i n g. A l s o, it a p p e a rs to us t h a t, a l t h o u gh t h e re is a g r e at d e al of s p e c i f i c i ty in t he g e n e t ic m a ke up and r e s p o n s e, s t i ll t he c o m p o u n ds t h at a re p r o d u c ed in r e s p o n se to i n f e c t i on a re not h i g h ly s p e c i f i c. A d i f f e r e nt p h y t o a l e x in is not n e e d ed f or e a ch i n f e c t i o us a g e n t. T he c e n t r al f a c t or b e c o m es w h e t h er t he p h y t o a l e x i ns a re p r o d u c ed s o on e n o u gh and in s u f f i c i e nt m a g- nitude. r e s i s t a n ce m e c h a n i s ms r e s i s t a n ce m e c h a n i s m s, a nd t he f a ct t h at r e s i s t a n ce a p p e a rs b a s ed on t he e x p r e s s i on of g e n e t ic i n f o r m a t i on s o on e n o u gh and in s u f f i c i e nt m a g n i t u d e. I hope I ' ve e m p h a s i z ed t he s i m i l a r i ty b e t w e en d i s e a se in p l a n ts and a n i m a l s, how we c an In o t h er w o r d s, i n d u ce We a re v e ry i n t e r e s t ed in a p p l y i ng t h is i n f o r m a t i on to p r a c- t i c al d i s e a se c o n t r o l. And w e ' ve e x t e n d ed t h e se t h o u g h ts to o t h er d i s e a s e s. W e ' ve w o r k ed w i th H e l m i n t h o s p o r i um d i s e a s es of c o r n, f i re blight of p e a rs and a p p l e s, a p p le s c a b, c o f f ee r u s t, c u c u m b er s c a b, and F u s a r i um wilt of m u s k m e l o n. It would be t r u ly e x c i t i ng to us and m o st r e w a r d i ng if we could i n d u ce r e s i s t a n ce in e m- b r y o n ic t i s s ue and c e l ls in t i s s ue c u l t u r e. P l a n ts d e v e l o p i ng f r om t h e se t i s s u es or c e l ls m ay h a ve t he m e t a b o l ic s w i t ch f or t a n ce p e r m a n e n t ly t u r n ed on. T he g e n e t i c i s ts of t he f u t u re m ay d e r i ve t h e ir r e s i s t a nt p l a nt s t o c ks f r om u n d i f f e r e n t i a t ed plant c e l ls in a t e st r e s i s- t u b e. D i s c u s s i on P e r i od D r. Duich; If I f o l l ow you c o r r e c t l y, c an we a s s u me t h at in a ny n a t u r al p l a nt c o m m u n i t y, w h e re we h a ve t he p o t e n t i al p r e s e n ce of m a ny s a p r o p h y t e s, and o b l i g a te and f a c u l t a t i ve p a r a s i t e s, we m i g ht get a n a t u r al i m m u n i z a t i on w h i ch could t r i g g er t he a c- c u m u l a t i on of t e r p e n o i d s. get a l a ck of a d i s e a se e p h i p h o t o t ic c o n d i t i o n. In m a ny s i t u a t i o n s, t h e r e f o r e, we m ay D r. K u c: I t h i nk t h is is t r u e. I!m p o i n t i ng at s o m e t h i ng e l s e. We f i nd t h at in c e r- D r. D u i c h: t a in a s p e c ts of t u r f, e s p e c i a l ly in b e n t g r a ss c u l t u r e s, an o b s e r v a- t i on t h at s e e ms to t ie in v e ry w e ll w h e re we m a ke a h i gh u se of f u n g i c i d e s. T h is i n v o l v ed t he u se of g e n e r al t y pe " c o n t a c t" f u n g i c i d es to b e g in with, and p a r t i c u l a r ly w h e re we h a ve c o n c e n- t r a t ed with i n d i v i d u al u se f u n g i c i d es and a r b i t r a r i ly s p r a y ed one s t r ip of bent c u l t i v a rs w i th one f u n g i c i de and a n o t h er s t r ip w i th a s e c o nd t y p e. L a te in t he s e a s o n, p a r t i c u l a r ly w i th p r e v e n t a t i ve t y pe s p r a y i n g, we o f t en s ee a c o m p l e te b r e a k d o wn of t he e f f e c t i v e- n e ss of t he f u n g i c i d e s. We n ow s e em to be s e e i ng t h is p r o b l em e v en m o re s e v e r ly with t he u se of s y s t e m ic f u n g i c i d e s. F or e x a m p l e, w e ' ve r e c o g n i z ed f or m a ny y e a rs t h at M e r i on b l u e- g r a ss h as a v e ry high d e g r ee of n a t u r al r e s i s t a n ce to H e l m i n- t h o s p o r i um v a g a ns in t he s p r i n g. T he o r g a n i sm d o es a t t a c k, but it m o re or l e ss is r e s t r i c t ed to a point. W e ' ve a l w a ys c o n s i d e r ed t h is to be a good r e a c t i on of n a t u r al r e s i s t a n c e, but w h en we u se t he s y s t e m ic f u n g i c i d e s, we s u d d e n ly f i nd t h at v a r i e t i es or c u l- t i v a rs w h i ch f or y e a rs and y e a rs h as b e en r e s i s t a nt to a p a t h o g e n, a re a ll of a s u d d en v e ry s u s c e p t i b l e. W e ' ve t a l k ed about s o me I d o n 't want to m i s l e ad you. D r. K u c: p h e n o ls and t e r p e n o i d s. D e p e n d i ng on t he p l a n ts u s ed in o ur s t u d i e s, d i f f e r e nt c o m p o u n ds a c c u m u l a t e. m ay be t e r p e n o i d s; in s o me c a s es c o u m a r i n s; in s o me c a s es s t e r o id g l y c o a l k y l o i d s; in s o me c a s es p h e n o ls like c h l o r o g e n i c, c a f f e ic a c id a re p a rt of t he d e f e n se m e c h a n i s m. i n t e r- a c t i o n s, s e v e r al of t h e se c o m p o u n ds m ay be p r o d u c e d. Not a ll In s o me c a s e s, t h e re In s o me o r g a n i s ms w i ll i n d u ce r e s i s t a n c e, and I d o n!t b e l i e ve one c o m- pound a l o ne e x p l a i ns r e s i s t a n c e. V e ry o f t e n, if w e ' re p r o t e c t i ng a g a i n st a C o l l e t o t r i c h u m, we u se d i f f e r e nt C o l l e t o t r i c h um spp. t h at d o n 't a t t a ck t h at v a r i e t y. With s o me o r g a n i s m s, w e ' ve had v e ry good luck. T h e y ' ve p r o t e c t ed a g a i n st a v e ry b r o ad s p e c t r um of f u n g i; f or e x a m p l e, A l t e r n a r i a, and d i f f e r e nt H e l m i n t h o s p o r i a. I t 's not u n r e a s o n a b le to t h i nk t h at in t he s o il i m m u n i z a t i on a g a i n st i m m u n i z a- p a t h o g e ns m ay o c c u r. W h en t h is b a l a n ce is d i s r u p t e d, t i on d o es not o c c u r. H o w e v e r, t h e re a re s e v e r al f a c t o rs i n v o l v ed w i th i n d u c ed p r o t e c t i o n. One, not a ll o r g a n i s ms w i ll p r o t e c t. S a p r o p h y t e s, a re s o me o r g a n i s ms t h at w o n 't g e r m i n a te on a p l a nt or g r ow a nd p e n e t r a te into a p l a n t. in g e n e r a l, a re not e f f i c i e nt in p r o t e c t i o n. T h e re D r. W h a l l e y: O n ce i n d u c e d, how long d o es t h is r e s i s t a n ce l a s t? D r. K u c: T he r e s i s t a n ce l a s ts t he l i fe of t he p l a nt in t he a r ea t h at y o u ' ve i n d u c ed it. T he d r a w b a c k, at l e a st with t he s y s t e ms t h at w e ' ve u s e d, is t h at we i n d u ce r e s i s t a n ce in t he a r ea we t r e a t. c o t yl w i ll be r e s i s t a nt f or t he l i fe of t he p l a n t. N ew g r o w th is not p r o t e c t ed u n l e ss we s p r ay it. We hope t h at we c an get a r o u nd t h is p r o b l e m. We c an p r o t e ct t i s s ue at a d i s t a n ce of about 0 . 5- 1.0 cm f r om t he i n d u c t i on s i t e. If we t r e at a s e c t i on of h y p o c o t y l, t h at s e c t i on of h y p o- t h at D r. W h a l l e y: Do you v i s u a l i ze a ny p o s s i b i l i ty of s p r a y i ng c r o ps a r t i f i c a l ly with an i n c o m p a t i b le i n o c u l um to p r o t e ct t h em f r om t he d e v e l o p m e nt of d i s e a s e? in c o n t r o l l i ng wilt In S o u t h e rn I n d i a n a, w a t e r m e l o ns a re n o r m a l ly i n o c u l um D r. K u c: C o o p e r a t i ng with t he H o r t i c u l t u re D e p a r t m e n t, we h a ve d o ne t h is on a p r a c t i c al s c a le f or t wo y e a rs of w a t e r m e l o n s. t r a n s p l a n t ed into t he f i e l d. of a n o n - p a t h o g en of w a t e r m e l on b e f o re t r a n s p l a n t i n g. T h is t e c h n i q ue g a ve us good r e s u l t s. S u r v i v al w as i n c r e a s ed and t he h o r t i c u l t u r i s ts w e re p l e a s ed with t wo y e a rs of t r i a l s. it is v e ry p r o m i s i ng to u se i n o c u l um w h i ch w i ll p r o t e c t. t h i nk i t 's e v en m o re p r o m i s i ng to u se s o me of t he c h e m i c al f a c- t o rs w h i ch w i ll i n d u ce I t 's no p r o b l em to dip in an I t h i nk I r e s i s t a n c e. D r. D a n i e l: E a r l i er you s a id r e s i s t a n ce j u st m o v ed up t he s t em a s m a ll a m o u n t, and n ow y o u ' re s a y i ng you t a ke a t r a n s p l a nt and dip it, as if it a f f e c t ed t he p l a nt f or a l o n g er p e r i o d. D r. Kuc: No, t he wilt is a r o ot p a t h o g e n. T h is wilt is an o r g a n i sm that g e ts into t he r o o t s. W h en we d ip t he r o ot s y s t em into t he s u s p e n s i o n, only t he t r e a t ed r o o ts a re p r o t e c t e d. D r. D a n i e l: T h e re a re n ew r o o t s? D r. Kuc; Y e s, n ew r o o ts a re f o r m e d. But a p p a r e n t ly t h e re enough of an e f f e ct t h e re to p r o t e ct it f or a long e n o u gh t i me get o v er a c r i t i c al p e r i o d. is to D r. D a n i e l: Out in n a t u re we o f t en s ee a d i s e a se a t t a ck b a r e ly s t a r t, t h en t he w e a t h er c h a n g e s. D o es a s l i g ht a t t a ck p r o t e ct a g a i n st t he next a t t a c k? it m a y. S i n ce t e m- t h e se I t h i nk u n d er s o me c i r c u m s t a n c es D r. K u c: a re b i o c h e m i c al p h e n o m e n a, t h ey w i ll be i n f l u e n c ed by S o m e- p e r a t u re and o t h er f a c t o rs t h at a f f e ct m e t a b o l ic s y s t e m s. t i m e s, j u st a s t r e ss c an i n d u ce r e s i s t a n c e. We c an i n d u ce t he a c c u m u l a t i on of p h y t o a l e x i ns by a p p l y i ng s t r e s s. By s t r e s s, I m e an s o me f u n g i c i d es or s o me a n t i m e t a b o l i t e s, w h i ch u n d er h i g h er l e v e ls would be t o x i c, but u n d er low l e v e ls w i ll p r o d u ce a s t r e ss and t r i g g er a s e r i es of e v e n ts w h i ch w i ll l e ad to t he a c c u m u- l a t i on of p h y t o a l e x i n s. M r. M a y e r: Along t h e se s a me l i n e s, you t a ke a v a r i e ty and it b e c o m es i n f e c t ed with a d i s e a se and m a y be 9 5% of it is w i p ed out, with a f ew r e m a i n i ng p l a n ts l e f t, t h en m a ke a s e l e c t i on out of t h is b e c a u se you a s s u me t h e se a re r e s i s t a n t. But a g a in t he d i s e a se a t t a c ks p r o g e ny of t h e se p l a n t s. Do you t h i nk t h at m a y be we h a ve t h is t r i g g e r i ng e f f e ct t he f i r st t i me and it d i d n 't c a r ry o v er i n to t he p r o g e n y? D r. K u c: T h a t 's p o s s i b l e. H o w e v e r, t he i n d u c i ng f a c t or and a g e n e t i c a l ly c o n t r o l l ed r e c e p t or s i te a re n e c e s s a ry f or p r o- t e c t i o n. W h en c r o s s es a re m a de and a s e g r e g a t i ng p o p u l a t i on is o b t a i n e d, d i f f e r e nt g e n e t ic c o m p l e m e n ts a re i n h e r i t ed and d i f f e r e nt r e s p o n s es to p a t h o g e ns a re o b s e r v e d. D r. P a y n e: You r e p o r t ed s p r a y i ng t he m a t e r i al onto m a t u re p l a n t s. H a ve you d o ne t h is and w as it e f f e c t i ve in i m m u n i z i n g? D r. K u c: Y e s. Now a g a in a w o rd of c a u t i o n. We h a ve not b e en s u c c e s s f ul in p r o t e c t i ng a g a i n st a r o ot p a t h o g en by s p r a y i ng t he f o l i a g e. W e ' ve only p r o t e c t ed t he t i s s ue t h at w e ' ve t r e a t e d, but we c an s p r ay m a t e r i a ls on p l a n ts and o b s e r ve p r o t e c t i o n. T E C H N I Q U ES F OR ASSESSING S E E D L I NG VIGOR AND E S T A B L I S H M E NT AND P O T E N T I AL A P P L I C A T I ON IN B R E E D I NG P R O G R A MS IN GRASSES R. D e r w yn B. W h a l l ey a nd C y r us M. M c K e ll l/ I n t r o d u c t i on D i f f e r e n c es in s u c c e ss of e s t a b l i s h m e nt a m o ng g r a ss s p e c i es r e s- a re w e ll k n o wn and h a ve b e en a t t r i b u t ed to d i f f e r e n t i al p l a nt p o n se to s u ch e n v i r o n m e n t al c o n d i t i o ns as t he p e r m e a b i l i ty of t he s o il to s e e d l i ng r o o t s, s o il m o i s t u re c o n t e n t, low or h i gh t e m p e r a- t u r e, and light i n t e n s i t y. S p e c i es t h at c o n s i s t e n t ly s h ow a r a p id g e r m i n a t i on r a t e, f a st r a t es of r o ot and t op g r o w t h, a r o b u st g r o w th h a b i t, or r e s i s t a n ce to s t r e ss a re o f t en r e f e r r ed to as h a v i ng s e e d l i ng v i g o r. S u c c e ss in s e e d l i ng e s t a b l i s h m e nt m ay be e n- h a n c e d, t h e r e f o r e, by e i t h er p r o v i d i ng f a v o r a b le e n v i r o n m e n t al c o n d i t i o ns or by s e l e c t i ng s p e c i es t h at h a ve a high d e g r ee of v i g or d u r i ng t he s e e d l i ng s t a g e. G r o w th of a n ew g r a ss p l a nt s t a r ts w i th a c t i v a t i on ( M a y er and P o l j a k o f f - M a y b e r, 1963) of t he e m b r yo and o t h er s e ed p a r t s, p r o c e e ds t h r o u gh t he s e e d l i ng s t a ge to t he p e r i od of r a p id v e g e t a t i ve g r o w th and, f i n a l l y, m a t u r i ty of t he e s t a b l i s h ed p l a nt ( W i l l i a m s, 1964) ( F i g. 1). With t he e m e r g e n ce of t he s e e d l i ng e s t a b l i s h m e nt b e g i ns and m ay not be c o n s i d e r ed a s u c c e ss u n t il t he plant h as d e v e l o p ed an a d e q u a te r o ot s y s t em and leaf a r ea to s u s t a in a h i gh r a te of g r o w th ( F i g. 2). W h a l l ey et al. ( 1966a) c o n s i d er t he s e e d l i ng s t a ge in t h r e e - p h a s e s: ( 1) t he h e t e r o t r o- phic s t a ge w h i ch o c c u rs f r om i n h i b i t i on to t he i n i t i a t i on of p h o- t o s y n t h e s i s; ling o b t a i ns c o m p l ex o r g a n ic c o m p o u n ds f r om both p h o t o s y n t h e s is t he t r a n s i t i on s t a ge d u r i ng w h i ch t i me t he s e e d- r a d i c l e, (2) l/ D e p a r t m e nt of B o t a n y, U n i v e r s i ty of N ew E n g l a n d, A r m i d a l e, N.S.W A u s t r a l i a, S t a te U n i v e r s i t y, L o g a n, U t ah 84321. 2351, and D e p a r t m e nt of R a n ge S c i e n c e, U t ah F i g u re 1. R e l a t i ve g r o w th c u r ve of b a r l e y. A d a p t ed f r om d a ta of R. F. W i l l i a ms ( 1964). T i me F i g u re 2. R e l a t i ve g r o w th c u r ve of b a r l e y. A d a p t ed f r om d a ta of R. F. W i l l i a ms ( 1964). T i me and t he r e m a i n d er of t he e n d o s p e r m, and (3) t he a u t o t r o p h ic s t a ge w h i ch is c o m p l e t e ly d e p e n d e nt on i ts own p h o t s y n t h e t ic p r o d u c t s. D u r i ng t he e s t a b l i s h m e nt s t a g e, i n t e n se c o m p e t i t i on f r om w i t h in t he s p e c i es and f r om o t h er s p e c i es m ay r e s u lt in t he l o ss of m a ny s e e d l i n g s, e v en u n d er f a v o r a b le e n v i r o n m e n t al c o n d i t i o n s. T h e re is no a p r i o ri v i g o r o us d u r i ng t he h e t e r o t r o p h ic s t a ge s h o u ld be e q u a l ly so d u r i ng t he a u t o t r o p h ic s t a g e, a p a rt f r om t he a d v a n t a g es b e s t o w ed by a r a p id s t a r t. With s u ch e x c e p t i o ns as t he c e r e a l s, m e m b e rs of t he G r a m i n e ae do not h a ve l a r ge s e e ds w i th t he a t t e n d a nt a d v a n t a ge of e x t e n s i ve f o od r e s e r v es a n d, as a r e s u l t, h a ve e v o l v ed v a r i o us s e e d l i ng c h a r a c t e r i s t i cs w h i ch a id in e s t a b l i s h m e n t. r e a s on why an i n d i v i d u al w h i ch is e f f i c i e nt and S e e d l i ng V i g or T h e re a re t wo d i s t i n ct a p p r o a c h es in d i s c u s s i ng s e e d l i ng v i g or in t he l i t e r a t u r e. T he a p p r o a ch u s ed by s e ed t e s t e rs w h e re t he o b j e c t i ve is to d e t e ct and u n d e r s t a nd r e d u c t i o ns r e l a t ed to a d v e r se c o n d i t i o ns of s e ed h a r v e s t, t r e a t m e nt or s t o r a ge ( H e y d e c k e r, 19 69a). And t he a g r o n o m ic a p p r o a ch w h e re t he o b- j e c t i ve is to u n d e r s t a nd and i m p r o ve s p e c i es a nd s t r a i ns with i n h e r e n t ly p o or s e e d l i ng v i g o r. in s e e d l i ng v i g or T he g e n e r al a im in a ll s e e d l i ng v i g or t e s t i ng is f or q u i c k, c o n v e n i e nt t e s ts w h i ch h a ve p r e d i c t i ve v a l ue c o n c e r n i ng t he f i e ld p e r f o r m a n ce of t he s e ed s a m p le t e s t e d. S e ed T e s t i ng A p p r o a ch T he n e c e s s i ty f or v i g or t e s t i n g, as d i s t i n ct f r om g e r m i n a t i on l o ts t e s t i n g, a r i s es b e c a u se t he f i e ld p e r f o r m a n ce of d i f f e r e nt s e ed m ay r a n ge f r om e x c e l l e nt to p o or d e s p i te s i m i l ar r e s u l ts f r om s t a n d a rd g e r m i n a t i on t e s ts ( P e r r y, 1967). T he V i g or T e st C o m- m i t t ee of t he I n t e r n a t i o n al S e ed T e s t i ng A s s o c i a t i on ( H e y d e c k e r, 19 69b) a t t e m p t ed to c o m p a re a n u m b er of v i g or t e s ts u s i ng t h r ee c a r e f u l ly s e l e c t ed l o ts of p ea and s ix of w h e at s e e d s. A n u m b er of c o o p e r a t i ng l a b o r a t o r i es p e r f o r m ed s e v e r al s t a n d a rd v i g or T he s t r i k i ng t h i ng a b o ut t he r e s u l ts w as t he h i gh c o r r e l a t i on b e- t w e en d i f f e r e nt t e s ts in r a n k i ng t he q u a l i ty of t he d i f f e r e nt s e ed t e s t s. l o t s. A g r o n o m ic A p p r o a ch M o st b i o l o g i s ts u se t he t e rm " s e e d l i ng v i g o r" in i ts b r o a d e st s e n se to d e s c r i be a v i g o r o us g r o w th h a b it t h at i n v o l v es a m o re r a p id s i ze i n c r e se t h an t h at of c o m p e t i ng p l a n ts of t he s a me age. P r o m pt s e e d l i ng g r o w t h, as s o on as c o n d i t i o ns b e c o me f a v o r a b l e, is an o b v i o us a d v a n t a ge t o w a rd s u c c e s s f ul e s t a b l i s h m e n t. l i n gs w i th a f a st r o ot and t op g r o w th m u st be s u p p o r t ed by e f f i c i e nt e n z y me s y s t e ms w i th w h i ch to m o b i l i ze s t o r ed f o od r e s e r v es in t he e n d o s p e rm t i s s u e. R a p id t r a n s p o rt of s o l u b le s u g a rs to g r o w i ng p o i n ts and a r e as of high m e t a b o l ic d e m a nd is a f u r t h er n e c e s s i ty f or a v i g o r o us s e e d l i n g. S e e d- S e e d l i ng v i g or is of c r i t i c al i m p o r t a n ce in p l a nt c o m p e t i t i o n. T he t r a d i t i o n al v i ew of C l e m e n ts ( 19 70) and l a t er of D o n a ld ( 19 63) h o l ds t h at c o m p e t i t i on is a p h y s i c al p r o c e ss of p l a n ts d r a w i ng f r om a p o ol of s u ch f a c t o rs as a i r, light, n u t r i e n ts and w a t e r. When t he p o ol is l i m i t ed or s u b j e ct to d e p l e t i o n, t h en t he m o re v i g o r o us s e e d l i ng t h at d r a ws r a p i d ly f r om t he p o ol or c an e x p l o re a g r e a t er v o l u me of t he e n v i r o n m e nt w i ll be t he s u c c e s s f ul c o m p e t i t o r. T e s ts f or s e e d l i ng v i g or h a ve b e en p r o p o s ed by n u m e r o us a u t h or as a m e a ns of e v a l u a t i ng or q u a n t i f y i ng c e r t a in a s p e c ts of s e e d l i ng g r o w t h. T he p r e d i c t i ve v a l ue of t h e se t e s ts in t e r ms of l a t er f i e ld p e r f o r m a n ce h as o f t en not b e en a d e q u a t e ly e s t a b l i s h e d. S o me of t he m o re r e c e nt w o r k e rs h a ve a t t a c k ed t h is p r o b l em a nd V a n d e r b i l t, 19 72; E d je and B u r r i s, 19 71; M o u t r ay and F r a k e s, 1971; W r i g ht and M a j o r, 1971) with v a r i a b le ( A b d u l l a hi s u c c e s s. C h a r a c t e r i s t i cs I m p o r t a nt in S e e d l i ng V i g or S e ed S i ze and Weight S e ed s i ze and w e i g ht a re e x t r e m e ly i m p o r t a n t. D a v i es ( 1927) s u m m a r i z ed r e s u l ts w i th c o m m o n ly u s ed B r i t i sh g r a s s es t h a t, " U n d er n o r m al f i e ld c o n d i t i o n s, t he s i ze of t he e n d o s p e rm is an i m p o r t a n t - f a c t or in d e t e r m i n i ng t he p o t e n t i al a b i l i ty of a s p e c i es to e s t a b l i sh i t s e l f ." C o n s i d e r a b le v a r i a t i on in s e ed s i ze s t a t i ng ( 19 66c). T h ey a l so r e p o r t ed t h at c o m m e r c i a l ly h a r v e s t ed e x i s ts in s e e ds p r o d u c ed f r om t he s a me p l a n t, a c c o r d i ng to W h a l l ey et al. s e e ds of P h a l a r is t u b e r o sa v a r. s t e n o p t e ra m ay c o n t a in a high p r o p o r t i on of i m m a t u re s e e d s. G e r m i n a t i on p e r c e n t a ge and r a te w e re l o w er in s m a ll and i m m a t u re s e e ds t h an in l a r ge m a t u re o n e s. W i t h in s p e c i e s, s e ed weight is of g r e a t er s i g n i f i c a n ce to s e e d l i ng v i g or t h an it is b e t w e en s p e c i es or g e n e r a. K i t t o ck and P a t t e r s on ( 1962) c o n c l u d ed t h a t, " T he c l o s er t he g e n e t ic b a c k- g r o u nd of c o m p a r ed l i n e s, t he h i g h er t he c o r r e l a t i on b e t w e en s e ed weight and s e e d l i ng v i g o r ." T h is g e n e r al r e l a t i o n s h ip h o l ds e v en down to t he i n d i v i d u al plant ( C a r l e t on and C o o p e r, 19 72). F i g u re 3. S e ed w e i g ht in r e l a t i on to t o t al s e e d l i ng l e n g t h. S c h i s m us a r a b i c us and O r y z o p s is m i l i a c ea s h ow a g r e a t er d e g r ee of c o r r e l a t i on b e t w e en s e ed weight and s e e d l i ng l e n g th t h an do t he v a r i o us P h a l a r is s p e c i es and v a r i e t i e s. In one of t he e a r l i e st p a p e rs p u b l i s h ed in t he J o u r n al of t he A m e r i c an S o c i e ty of A g r o n o m y, Z a v i tz ( 1908) r e p o r t ed c o n s i s- t e n t ly g r e a t er y i e l ds f r om p l a n t i n gs of l a r ge w e l l - f i l l ed c e r e al s e e ds t h an f r om s m a ll w e l l - f i l l ed s e e d s. M a ny r e p o r ts e m p h a s i ze t he i m p o r t a n ce of a g r e a t er v o l u me of e n d o s p e rm food r e s e r ve in r e l a t i on to s u b s e q u e nt s e e d l i ng s i ze or r a te of g r o w th (Hove and K l e i n e n d o r s t, 1962; D e m i v l i c a k m ak et a l ., 1963; K n e e b o ne and C r e m e r, 1955). W h a l l ey et al. ( 1966a) s h o w ed t h at l a r ge s e e ds h ad l o n g er s e e d l i ng g r o w th t h an s m a ll s e e ds of t he s a me s p e c i es ( F i g. 3). One e x t r e m e ly s m a ll s e e d ed s p e c i e s, S c h i s m us g r ew at a f a st r a te but, in t he a b s e n ce of light, w as t h r o u gh g r o w- ing e v en b e f o re r a d i c le e m e r g e n ce of l a r g e r - s e e d ed s p e c i es had s t a r t e d^ a r a b i c u s, S e ed s i ze and w e i g ht a re not as i m p o r t a nt as t he p l a nt r e s- p o n s es d e p e n d e nt on t h e m, s u ch as e m e r g e n ce f r om s o il d e p t h s, In a r e as s u b j e ct to e a r ly s e e d l i ng r o ot g r o w th and e m b r yo s i z e. r a p id d r y i ng of t he s o il s u r f a c e, d e e p er p l a n t i ng of g r a ss s e e ds is i m p o r t a nt f or s u c c e s s f ul e s t a b l i s h m e nt and s u r v i v a l. R o g l er ( 19 54) and V o g el ( 19 63), w o r k i ng with A g r o p y r on c l o se c o r r e l a t i on b e t w e en s e ed w e i g ht and t o t al e m e r g e n ce f r om d e p t hs g r e a t er t h an one i n c h. c r e a s ed t he r a te of e m e r g e n c e. S i m i l ar r e s u l ts w e re r e p o r t ed by M u l t a m a ki ( 19 62) and K a l t on et al. ( 19 59). I n c r e a s i ng t he s e e d i ng d e p th d e- s p p ., r e p o r t ed a E a r ly s e e d l i ng r o ot g r o w th is a l m o st e q u a l ly i m p o r t a nt as e m e r g e n ce and h as b e en s h o wn to be c o r r e l a t ed w i th s e ed s i ze ( P l u m m e r, 1943; K i t t o ck a nd P a t t e r s o n, 1962; T a d m o re and Cohen, r e- 19 68). E a r ly r o ot g r o w th is l a r g e ly d e p e n d e nt on e n d o s p e rm s e r v es b e c a u se p h o t o s y n t h e t ic a c t i v i ty of l e a v es is d e l a y ed a f t er e m e r g e n ce ( A n s l o w, 19 62) and is i n i t i a l ly i n s u f f i c i e nt to s u p p o rt r o ot g r o w th and m e t a b o l i s m. R e c u r r e nt s e l e c t i on f or i n c r e a s ed s e ed s i ze in at l e a st two p e r e n n i al g r a ss s p e c i e s, P a n i c um a n t i d ó t a le ( W r i g ht and M a j o r, 19 71) and B r o m us i n e r m is ( T r u pp and C a r l s o n, 19 71), h as l e ad to i n c r e a s ed s e e d l i ng v i g o r, as m e a s u r ed by s e v e r al c r i t e r i a. T h e re would p r o b a b ly be w i de v a r i a b i l i ty in t he p o s s i b le r a te of p r o g r e ss in both s e ed s i ze a nd s e e d l i ng v i g or w i th d i f f e r e nt g r a ss s p e c i e s. Rapid G e r m i n a t i on R a p id g e r m i n a t i on is a n o t h er c h a r a c t e r i s t ic t h at m ay c o n t r i- bute to a v i g o r o us and s u c c e s s f ul s e e d l i n g. S e e ds w h i ch g e r m i n- ate r a p i d ly h a ve o b v i o us a d v a n t a g es p h y s i c al f a c t o rs of t he e n v i r o n m e n t. With a r a p i d ly d r y i ng s e e d- bed, t h is a d v a n t a ge b e c o m es e v en m o re i m p o r t a n t. H o r n i ng and Canode ( 19 63) u s ed a v i g or s c o re s i m i l ar to t he g e r m i n a t i on s p e ed index of M a g u i re ( 1962) to e v a l u a te s m o o th b r o m e g r a ss s e ed h a r- v e s t ed u n d er d i f f e r e nt c o n d i t i o n s. T h ey f o u nd a good c o r r e l a t i on b e t w e en t he v i g or s c o re a nd o t h er m e t h o ds of t e s t i ng s e ed q u a l i t y. in t e r ms of c o m p e t i t i on f or R a p id g e r m i n a t i on is o f t en a c h a r a c t e r i s t ic of w e e dy g r a s s es that i n v a de g r a ss c o m m u n i t i es w h en a s h o rt p e r i od of f a v o r a b le conditions o c c u rs ( M a j or et a l ., c h e m i c al s y s t e ms t h at a re a c t i ve in t he g e r m i n a t i on p r o c e ss c o n- tinue to o p e r a te in t he s e e d l i ng s t a ge and e v en b e y o n d, t h us taining a v i g o r o us g r o w th h a b i t. i960). T he s a me i n t e r n al b i o- s u s- B i o c h e m i c al and P h y s i o l o g i c al A c t i v i ty A h i gh l e v el of b i o c h e m i c al and p h y s i o l o g i c al a c t i v i ty of s e e d l i n gs is b a s ic to s e e d l i ng v i g o r. S e e d l i n gs w i th h i gh p h y- s i o l o g i c al a c t i v i ty g e n e r a l ly a re t h o se with a h i gh r a t i ng f or v i g o r. M a y er and P o l j a k o f f - M a y b er ( 1963) d i s c u ss t he m a ny s y s t e ms and p r o c e s s es t h at m ay be i n v o l v ed in s u p p o r t i ng s e e d l i ng g r o w t h. A f t er i m b i b i t i o n, s t o r a ge p r o d u c ts a re e n z y m a t i c a l ly c h a n g ed to s o l u b le f o r m s; m e t a b o l i sm a nd f u r t h er b r e a k d o wn of s t o r a ge p r o d u c ts s u ch as c a r b o h y d r a t e s, containing c o m p o u n ds i n c r e a s es r e s u l t i ng in h i gh d e m a n ds f or e n e r g y - r i ch s u b s t r a te and o x y g e n. C e ll d i v i s i on and e l o n g a t i on growing p o i n ts a re h i g h ly d e p e n d e nt on e f f i c i e nt t r a n s p o rt in o r d er to s u s t a in h i gh g r o w th l i p i d s, p r o t e i n s, and p h o s p h o r u s- in s y s t e ms r a t e s. R e p o r ts on m e t a b o l ic r a te as an i n d ex of p h y s i o l o g i c al a c t i- vity in v i g o r o us s e e d l i n gs a re c o n f l i c t i n g. M a h a d e v a p pa ( 1967) r e p o r t ed t h at leaf t i s s u es of p e a rl m i l l et f a i l ed to s h ow any d i f- f e r e n c es in t he r a te of r e s p i r a t i o n, n or could he f i nd a c o r r e l a- tion b e t w e en r e s p i r a t i on c a p a c i ty and g r o w th r a t e. E a r l i e r, F i g u re 4 -A to t u b e r o sa C o n v e r s i on of s t a r ch s u g a rs of P h a l a r is v a r. s t e n o p t e ra S p r i n g s) w h i ch is l ow in s e e d l i ng v i g o r. ( T u le r a t ed F i g u re 4 -B to C o n v e r s i on of s t a r ch s u g a rs of P. c o e r u l e s c e ns w h i ch is r a t ed high in s e e d l i ng v i g o r. O 20 T h r o n e b e r ry and S m i th ( 19 55) r e p o r t ed e s s e n t i a l ly t he s a me sult with g e r m i n a t i ng c o rn s e e d s. H o w e v e r, ( 19 68), W o o d s t o ck and F e e l ey ( 19 65) and C a n t r e ll et al. r e p o rt c l o se c o r r e l a t i o ns b e t w e en r e s p i r a t i on r a t es and s e e d l i ng vigor. r e- K i t t o ck and L aw ( 19 72) R a p id m o b i l i z a t i on of e n d o s p e rm r e s e r v es and t r a n s p o rt a p p e a rs to be a n o t h er f a c t or in s e e d l i ng v i g o r. W h a l l ey et al. ( 19 66b) and W h a l l ey and M c K e ll ( 19 67) s h ow t h at t he d e g r ee of a v a i l a b i l i ty of s u g a rs or t he l a ck of s u f f i c i e nt t r a n s p o rt to t he growing s e e d l i ng m ay p a r t i a l ly a c c o u nt f or d i f f e r e n c es in s e e d- ling v i g or a m o ng s e v e r al s p e c i es of P h a l a r is ( F i g. 4). C o n s i d- e r a b le s t u dy is n e e d ed b e f o re a c l e ar p i c t u re c an be d r a wn of the r a te of b i o c h e m i c al and p h y s i o l o g i c al a c t i v i ty in v i g o r o us s e e d l i n g s. F o r ce of G r o w th F o r ce of s e e d l i ng g r o w th is a v a l u a b le p l a nt a t t r i b u te t h at m ay be c r i t i c al in s u r v i v a l. W h e re r a p i d i ty of e m e r g e n ce d e p e n ds on a high g r o w th r a te of t he c o l e o p t i le and s h o o t, t he f o r ce of e m e r g e n ce m ay be m o re d e p e n d e nt on e n d o s p e rm r e s e r v es and a d u r a b le e p i d e r m i s. K a l t on et al. ( 19 59) m a de o b s e r v a t i o ns on t he r e l a t i on b e t w e en s e ed s i ze and e m e r g e n ce f r om d i f f e r e nt d e p t h s. E m e r g e n ce w as d e l a y ed or r e d u c ed w i th an i n c r e a se in s e e d l i ng depth but t he d i f f e r e n ce in s e e d l i ng e m e r g e n ce b e t w e en l a r ge and s o w ed s e e ds of E l y m us s m a ll s e e ds i n c r e a s e d. L a w r e n ce ( 1963) j u n c e us at d e p t hs of f r om 0.5 to 1.5 in. and r e p o r t ed a p o s i t i ve c o r r e l a t i on b e t w e en t o t al e m e r g e n ce and s e ed s i z e. s e ed y i e ld and f o r a ge y i e ld w e re a l so c o r r e l a t e d. S u b s e q u e n t l y, R a te of G r o w th R a te of g r o w t h, b e i ng one of t he m o st o b v i o us c h a r a c t e r i s t i c s, is o f t en r e g a r d ed as e v i d e n ce of s e e d l i ng v i g o r. B o th r a p id g r o w th and r o ot g r o w th c an be a d i s t i n ct a d v a n t a ge in s e e d l i ng e s t a b l i s h m e n t. E a r ly a c h i e v e m e nt of an a d e q u a te leaf a r ea s u p- p l i es t he n e e ds of t he d e v e l o p i ng s e e d l i ng f or p r o d u c ts of p h o t o- s y n t h e s is in a d d i t i on to o c c u p y i ng s u r f a ce a r ea t h at would o t h e r- w i se be u s ed by c o m p e t i ng s p e c i e s. L o o m is a nd W i l l i a ms ( 19 63) t op leaf s u g g e st t h at t he t i me r e q u i r ed f or a t t a i n m e nt of an o p t i m um a r ea and, t h u s, i n c r e a s ed p r o d u c t i v i ty c an be s h o r t e n ed by p l a n t i ng m o re s e e ds p er unit a r ea or by p l a n t i ng s e e ds t h at p r o d u ce a l a r ge i n i t i al leaf a r ea p er p l a n t. T h o m as ( 1966) s h o w ed t h at leaf s i ze is c o r r e l a t ed w i th s e ed s i ze but t h at t he r e l a t i o n s h ip d e c r e a s es w i th t i m e. S h i b l es and M a c D o n a ld ( 19 62) c o m p a r ed t wo v a r i e t i es of b i r d s f o ot t r e f o il w h i ch w e re k n o wn f or low and high s e e d l i ng v i g o r. E v en t h o u gh both v a r i e t i es w e re e q u al in net p h o t o s y n t h e- t ic r a te p er unit a r ea of c o t y l e d o n, t he v a r i e ty k n o wn f or i ts high s e e d l i ng v i g or h ad a h i g h er r a te of p r o d u c t i on of photo s y n t h e t ic s u r f a c e, e v en t h o u gh s e e ds of t he s a me s i ze w e re u s e d. R o ot S y s t em D e v e l o p m e nt R a p id d e v e l o p m e nt of a r o ot s y s t em s u f f i c i e nt to g a r n er w a t er and n u t r i e n ts f or t he young s e e d l i ng is p e r h a ps a h i g h er p r i o r i ty t h an e a r ly t op g r o w t h, if e n d o s p e rm r e s e r v es l a r ge s e e ds a re s t i ll a v a i l a b l e. P l u m m er (1943) a t t r i b u t ed t he s u c c e ss or f a i l u re of e s t a b l i s h m e nt of t w e l ve r a n ge g r a s s es to t o t al r o ot d e v e l o p m e nt p r i or to s u m m er d r o u g h t. T he m o st s u c c e s s f ul s p e c i e s, A g r o p y r on c r i s t a t u m, p r o d u c ed a g r e a t er t o t al r o ot l e n g th in t he s e e d l i ng s t a g e. D i f f e r e n t i al r e s p o n se of t o ps and r o o ts to t e m p e r a t u re m u st be c o n s i d e r e d. B r o u w er ( 1966) r e- v i e w ed t he l i t e r a t u re c o n c e r n i ng f a c t o rs a f f e c t i ng r o ot g r o w th r a t es and p r e s e n t ed i n f o r m a t i on s h o w i ng t h at at low r o ot g r o w th is f a v o r ed m o re t h an t op g r o w t h. An i n c r e a se t e m p e r a t u re r e s u l ts in a b u r st of t op g r o w th giving an in s h o ot r o ot r a t i o. O t h er f a c t o rs s u ch as low n i t r o g e n, m o i s t u re s t r e s s, a nd s h a de a l so c a u se low s h o o t / r o ot i n c r e a se r a t i o s. in t e m p e r a t u r e s, in C o h en and T a d m or (19 69) r e p o r t ed l a r ge d i f f e r e n c es a m o ng 10 s p e c i es of g r a s s es and l e g u m es in s e e d l i ng r o ot g r o w t h. S e e d- ling r o ot e l o n g a t i on in t he u p p er s o il l a y er (2 to 12 c m) w as g r e a t- e st at 25 ° C. W h e r e as in t he d e e p er l a y er (12 to 22 c m ), 2 0 °C a p p e a r ed to be t he m o st f a v o r a b le t e m p e r a t u r e. S e ed s i ze a l so c o r r e l a t ed w e ll w i th r a te of r o ot e l o n g a t i o n. P h a l a r is t u b e r o sa s e e ds w e re t he s m a l l e st in t h e ir s t u dy and h ad t he s l o w e st e l o n g a- t i on r a t e, w h e r e as t he o p p o s i te w as t r ue f or w h e at and b a r l e y. L ow t e m p e r a t u r es a c c e n t u a t ed t h e se d i f f e r e n c e s. F or s p e c i es like P h a l a r i s, w h i ch is a d a p t ed to M e d i t e r r a n e an c l i m a t e s, s e ed v a r i e t i es m u st be d e v e l o p ed to p r o v i de i n c r e a s ed v i g or of r o ot g r o w th to o v e r c o me t he a d v e r se e f f e c ts of low s o il t e m p e r a- t u r es d u r i ng t he m o i st s e a s o n. H y d er et al. (1971) a l so a t t r i b u t ed d i f f e r e n c es in t he e a se of e s t a b l i s h m e nt b e t w e en b l ue g r a ma and c r e s t ed w h e at g r a ss to d i f f e r e n c es in r o o t i ng d e p t h. l a r g er R e s i s t a n ce to E n v i r o n m e nt R e s i s t a n ce to p h y s i c al and b i o l o g i c al e n v i r o n m e nt m ay be a d i r e ct e x p r e s s i on of s e e d l i ng v i g or c a u s ed by a t o u gh e p i d e r m i s, an a n a t o m i c al or p h y s i o l o g i c al m a k e up t h at m a k es a s e e d l i ng n o n- s u s c e p t i b le to p l a nt p a t h o g e n s, or a p h y s i c al c o n s t i t u t i on t h at e n a b l es a s e e d l i ng to w i t h s t a nd m e c h a n i c al d a m a ge of v a r i o us t y p e s. C o n- v e r s e l y, r e s i s t a n ce as an a s p e ct of s e e d l i ng v i g or m ay be m e r e ly t he r e s u lt of a high r a te of g r o w t h, t he a b i l i ty to g r ow at an u n u s u al t e m p e r a t u re r e g i m e, or t he a b i l i ty to w i t h s t a nd o t h er e n v i r o n- m e n t al s t r e s s es b e c a u se of a h i gh l e v el of b i o c h e m i c al or p h y s- i o g i c al a c t i v i t y. W h a t e v er t he c a s e, a r e s i s t a nt s e e d l i ng m ay e a s i ly be c l a s s ed as a v i g o r o us s e e d l i ng if it r e s u l ts in a s u b s t a n- t i al a d v a n t a ge in e s t a b l i s h m e n t. S e v e r al s t u d i es give e x a m p l es of t he s i g n i f i c a n ce of r e s i s t a n c e. H i gh t e m p e r a t u r es and h i gh h u m i d i ty p r i or to g e r m i n a t i on w e re s h o wn by H e l m e rs et al. ( 19 62) to e l i- m i n a te s e e ds t h at did not h a ve t he p o t e n t i al f or h i gh s e e d l i ng v i g o r. D i s e a se r e s i s t a n ce ( I s l e y, 1957) is a n o t h er a s p e ct of p l a nt v i g or but one w h i ch d r a ws on m o re f u n d a m e n t al s e e d l i ng c h a r a c t e r i s- t i c s, s u ch as a n a t o my and p h y s i o l o g y. R e s i s t a n ce of r e ed c a n a r y- g r a ss s e e d l i n gs to f l o o d i ng or a c id c o n d i t i o ns w as c o r r e l a t ed with f i e ld p e r f o r m a n ce ( M a rk and M c K e e, 1968), t h us i n d i c a t i ng i m p o r t a n ce of t h is t y pe of v i g or p l a nt p e r f o r m a n c e. in t he s e e d l i ng s t a ge f or o v e r a ll t he E n v i r o n m e n t al C o n d i t i o ns and S e e d l i ng G r o w th T e m p e r a t u re T h e re is w i de v a r i a b i l t iy in t he t e m p e r a t u re o p t i ma of d i f- in t he e a r ly and f e r e nt s p e c i es and o f t en d i f f e r e nt r e q u i r e m e n ts l a t er s t a g es of s e e d l i ng g r o w th ( C o h en and T a d m o r, 19 69). In m i x ed s t a n d s, d i f f e r e nt t e m p e r a t u re c o n d i t i o ns u s u a l ly f u n c t i on t h r o u gh c o m p e t i t i on in t h at a s p e c i es f a v o r ed by t he t e m p e r a t u re c o n d i t i o ns h as a c o m p e t i t i ve a d v a n t a ge o v er o t h er s p e c i e s. T he i m p o r t a n ce of t e m p e r a t u re a d a p t a t i on s h o u ld not be o v e r l o o k ed in b r e e d i ng p r o g r a m s. W a t er g e r m i n a te at 4 -6 b a rs b e l ow A g a i n, s p e c i es v a ry w i d e ly in t h e ir r e s p o n se to w a t er a v a i l- a b i l i ty d u r i ng g e r m i n a t i on and e s t a b l i s h m e n t. E a sy to e s t a b l i sh s p e c i e s, s u ch as a n n u al r y e g r a s s, w i ll t he l i m i t i ng w a t er p o t e n t i al of o t h er m o re d i f f i c u lt s p e c i es W i l l i am et a l ., 19 70; M c W i l l i am and P h i l l i p s, 19 71). T u r f g r a s s es a re o f t en e s t a b l i s h ed u n d er l e ss t h an i d e al s o il m o i s t u re c o n- d i t i o ns and, u n d o u b t e d l y, s e l e c t i on f or g e r m i n a t i on at low s o il w a t er p o t e n t i a ls would be v a l u a b l e. T h e re d o es not a p p e ar to be a ny r e l a t i o n s h ip b e t w e en g e r m i n a t i on at low w a t er p o t e n t i al and d r o u g ht t o l e r a n ce of t he m a t u re p l a n ts ( M c W i l l i am and P h i l l i p s, 1971). ( M c- F e r t i l i ty A c u l t u r al p r a c t i c e, s u ch as f e r t i l i z a t i o n, is an e a sy w ay of i n- c r e a s i ng t he c o m p e t i t i ve a d v a n t a ge of d e s i r a b le s p e c i e s, p r o v i d ed t he d i f f e r e n t i al r e s p o n s es of t he s p e c i es p r e s e nt a re known. L e ss w e ll u n d e r s t o od a re t he d i f f e r e nt r e q u i r e m e n ts of s p e c i es at v a r i- o us s t a g es d u r i ng s e ed g e r m i n a t i on and e s t a b l i s h m e n t. A y e ke and M c K e ll (19 69) s h o w ed t h at t he a d d i t i on of n i t r o g en with p h o s p h o r us d e c r e a s ed t he t op and r o ot g r o w th of O r y z o p s is m i l i a c ea s e e d l i n gs d u r i ng t he f i ve d a ys f r om r a d i c le e m e r g e n ce in t he d a r k. With L o l i um m u l t i f l o r u m, l o w er r a t es of N i n c r e a s ed g r o w th r a t es w h i ch t h en d e c l i n ed at h i g h er r a t e s. W h a l l ey and M c K e ll ( u n- p u b l i s h ed d a t a) h a ve r e c e n t ly s h o wn t h at c r e a s t ed w h e a t g r a ss a nd R u s s i an w i l d r ye s e e d l i n gs r e s p o nd to p h o s p h o r us d u r i ng t he e a r ly s t a g es of g r o w t h, and t h at a r e s p o n se to n i t r o g en d o es not b e c o me a p p a r e nt u n t il t wo w e e ks f o l l o w i ng s e e d l i ng e m e r g e n c e. r e s p o n s e s. T h e re w e re a l so s o me d i f f e r e n t i al s p e c i es and s t r a in T a b le 1. S e e d i ng r a t es a nd s e e d l i ng s u r v i v al of a d r y l a nd p e r e n n- i al g r a ss c o m p a r ed w i th a c o m m on t u r f g r a ss ( d a ta c a l c u l a t ed to c o m m on u n i ts and r o u n d ed off). r e- S p e c i es C r e s t ed W h e a t g r a ss K e n t u c ky B l u e g r a ss 56 g m / l 0 0 m2" l 5 00 g m / l 0 0 m2^ Seeding r a te No. of v i a b le s e e d s / g r am No. of v i a b le s e e ds s o w n / m2 No. of s e e d l i n gs 7 d a ys a f t er s o w i n g / m2 No. of s e e d l i n gs 9 d a ys a f t er s o w i n g / m2 No. of s e e d l i n gs 14 d a ys a f t er s o w i n g / m2 No. of p l a n ts 12 m o n t hs a f t er s o w i n g / m2 P e r c e n t a ge e s t a b l i s h m e nt 346+ 200 30''* 44'^ 25'^ 12 3,570# 54,000 11,000^ 20 "Hyder et al. 19 55, +W h y te et al. 19 59, # M u s s er a nd P e r k i n g s, 19 69 It is c l e ar t h at t he r e s p o n s es of g r a ss s e e d l i n gs to s o il n u t r i e nt l e v e ls d u r i ng t he e a r ly s t a g es of e s t a b l i s h m e nt a re l i t t le u n d e r s t o o d, but a re of g r e at i m p o r t a n ce f or t he e s t a b l i s h m e nt of t u r f g r a s s e s. U n d o u b t e d ly t h e re is s c o pe f or b r e e d i ng w o rk to s e l e ct s t r a i ns w h i ch c an r e s p o nd to h i gh f e r t i l i ty l e v e ls d u r i ng t he e s t a b l i s h m e nt p h a s e. T e c h n i q u es f or M e a s u r i ng S e e d l i ng E s t a b l i s h m e nt T he t e c h n i q u es f or m e a s u r i ng s e e d l i ng e s t a b l i s h m e nt h a ve not b e en s t u d i ed to t he s a me e x t e nt as s e e d l i ng v i g o r. H o w e v e r, t h e re a re p r o b l e ms i n v o l v e d, p a r t i c u l a r ly w i th r e f e r e n ce to t he d e f i n i t i on of s t a nd e s t a b l i s h m e nt and t he t i me of a s s e s s m e n t. T he end of s e e d l i ng e s t a b l i s h m e nt could be t a k en as t he end of t he lag p h a se w h en t he s e e d l i ng h as d e v e l o p ed a good r o ot s y s t em and is c o m m e n c i ng i ts p e r i od of r a p id g r o w th ( F i g. 2). F ew r e p o r ts a re a v a i l a b le w h e re r e p e a t ed s e e d l i ng c o u n ts h a ve b e en m a d e. E x a m p l es of s u ch d a ta a re g i v en in T a b le 1. If s e e d i ng r a t es s i m i l ar to t h o se u s ed f or l a wn g r a s s es a re u s ed in a d r y l a nd s i t u a t i o n, t he i n t r a s p e c i f ic c o m p e t i t i on is e x- t r e m e ly high, and t he i n d i v i d u al p l a n ts d e v e l op p o or r o ot b e c a u se of i n t e n se c o m p e t i t i o n. D u r i ng a p e r i od of s t r e s s, t he w h o le s t a nd is l i k e ly to s u c c u m b. A v a l id a s s e s s m e nt of s t a nd e s t a b l i s h m e nt c a n n ot be m a de u n t il p l a nt n u m b e rs a re m o re or l e ss c o n s t a nt with t i me a nd s h o u ld i n c l u de b e l ow as w e ll as a b o v e- g r o u nd i n f o r m a t i o n. T h e re is a n e ed f or b a s ic d a ta r e g a r d i ng t he s h a pe of t he s u r v i v o r s h ip c u r v es f or d i f f e r e nt s p e c i es u n d er d i f- f e r e nt s i t u a t i o n s. s y s t e ms P h i l o s o p hy of V i g or T e s t i ng In a ny e v a l u a t i on of t u r f g r a ss b r e e d i ng m a t e r i al f or s e e d- ling v i g o r, t he i n c l u s i on of s p e c i es a d a p t ed to e s t a b l i s h m e nt u n d er r a n ge c o n d i t i o ns is d e s i r a b l e. S u ch s t u d i es would p r o v i de v a l u a b le i n s i g h ts to t he p l a nt b r e e d er who w o r ks w i th s p e c i es i n t e n d ed f or l e ss d e m a n d i ng e n v i r o n m e n t s. In a d d i t i o n, t he e n v i r o n m e nt p r o- v i d ed by t he a v e r a ge a m a t e ur l e ss t h an l a wn g r o w er is u s u a l ly s o m e w h at i d e a l. T he s e q u e n ce of e v e n ts f r om i m b i b i t i on to m a t u r i ty is e s s e n- t i a l ly a c o n t i n u u m. F ar t oo o f t en p l a nt b r e e d e rs s e e d l i ng v i g or h a ve a s s e s s ed m a t e r i al at only one or t wo In no c a se h as t he p r e d i c t i ve v a l ue of a s i n g le s e e d l i ng v i g or t e st f or s e e d l i ng p e r f o r m a n ce d u r i ng t he e n t i re c o n t i n u um b e en s a t i s f a c t o r i ly e s t a b l i s h e d. S e v e r al i m p o r t a nt t e s ts a re i z ed b e l o w: i n t e r e s t ed s u m m a r- in t i m e s. G e r m i n a t i on U n d er W a t er S t r e ss ( M c W i l l i am a nd P h i l l i p s, 19 70) T he a b i l i ty to g e r m i n a te u n d er w a t er s t r e ss is u n d o u b t e d ly i m p o r t a nt in m a ny s i t u a t i o n s, and t he m e a s u r e m e n ts c an be p e r- f o r m ed on a r o u t i ne b a s i s. R a te of G e r m i n a t i on ( M a g u i r e, 1962) T he s p e ed of g e r m i n a t i on is e a s i ly m e a s u r ed u n d er s t a n d a rd c o n d i t i o ns a nd c an be an i m p o r t a nt c h a r a c t e r i s t ic u n d er a d v e r se e n v i r o n m e n t al c o n d i t i o n s. S e ed S i ze ( T r u pp and C a r l s o n, 19 71) S e l e c t i on f or s e ed s i ze h as b e en s h o wn to be c o r r e l a t ed w i th e a r ly g r o w th r a te and is e a s i ly m e a s u r e d. S e ed R e s p i r a t i on S t u d i es ( W o o d s t o c k, 19 69) T h e re a p p e a rs to be good c o r r e l a t i on b e t w e en e a r ly s e ed r a t e s. r e s p i r a t i on r a t e s, R Q v a l u es and e a r ly s e e d l i ng g r o w th H o w e v e r, it is e s s e n t i al to m e a s u re r e s p i r a t i on r a t es of i n d i v i- d u al s e e ds w h i ch p r e s e n ts t e c h n i c al d i f f i c u l t i es w i th t he s m a ll s e e d ed t u r f g r a s s e s. G r o w th M e a s u r e m e n ts ( H e y d e c k e r, 19 69a p. 206) T he m e a s u r e m e nt of t op and r o ot g r o w th of a m a ss of s e e d- l i n gs at s o me point in t i me a f t er s e e d i ng d o es not give t he i n f o r- m a t i on r e q u i r ed in v i ew of t he s m a ll p e r c e n t a ge of p l a n ts w h i ch s u r v i ve to m a t u r i t y. S u ch m e a s u r e m e n ts m u st be on an i n d i v i- d u al and r e p e a t ed b a s i s; t e d i o us f or p l a nt t o ps and d i f f i c u lt f or p l a nt r o o t s. F u r t h er i n f o r m a t i on on s u r v i v o r s h ip c u r v es in a t u r f g r a ss s i t u a t i on a re n e c e s s a r y. S e e d l i ng v i g or p r e s e n ts c o n s i d e r a b le p r o b l e ms to t he plant b r e e d er b e c a u se of t he m a ny f a c t o rs i n v o l v e d. N e v e r t h e l e s s, m o st p r o m i s i ng a p p r o a ch s e e ms to be one b a s ed on c o n t i n u al m u l t i a s s e s s m e nt of m a t e r i al on an i n d i v i d u al b a s is r a t h er r e l i a n ce on a ny s i n g le v i g or t e s t. t he t h an R e f e r e n c es C i t ed A b d u l l a h i, A., and R. L. V a n d e r b i l t. 1972. R e l a t i o n s h i ps of v i g or t e s ts and s e ed s o u r ce a nd s i ze to s o r g h um s e e d l i ng e s t a b- l i s h m e n t. A g r o n. J. 64: 143-144. A n s l o w, R. C. 19 62. A q u a n t i t a t i ve a n a l y s is of g e r m i n a t i on and e a r ly s e e d l i ng g r o w th in p e r e n n i al r y e g r a s s. J. B r i t. G r a s s- l a nd Soc. 17: 2 6 0 - 2 6 3. A y e k e, C. A., and C. M. M c K e l l. 1969. E a r ly s e e d l i ng g r o w th of I t a l i an r y e g r a ss and s m i lo as a f f e c t ed by n u t r i t i o n. M g m t. 22: 2 9 - 3 2. J. R a n ge B r o u w e r, R. 1966. 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M a d i s o n, W i s. pp. 4 7 4 - 4 9 0. P e r r y, D. A. 19 67. S e ed v i g o ur and f i e ld e s t a b l i s h m e nt of p e a s. P r o c. Int. S e ed T e st A s s. 32: 3-12. P l u m m e r, A. P. 1943. T he g e r m i n a t i on and e a r ly s e e d l i ng d e v e l- o p m e nt of t w e l ve r a n ge g r a s s e s. 35: 19-34. J. A m e r. Soc. A g r o n. R o g l e r, G. A. 19 54. S e ed s i ze and s e e d l i ng v i g or in c r e s t ed w h e a t. A g r o n. J. 46: 216-220. S h i b l e s, R. M., and H. A. M a c D o n a l d. 19 62. P h o t o s y n t h e t ic a r ea and r a te in r e l a t i on to s e e d l i ng v i g or of b i r d s f o ot t r e f o il ( L o t us c o r n i c u l a t u s ). C r op Sci. 2: 2 9 9 - 3 0 2. T a d m o r, N. H., and Y. C o h e n. 1968. P r e - e m e r g e n ce s e e d l i ng r o ot d e v e l o p m e nt of M e d i t e r r a n e an g r a s s es and l e g u m e s. C r op Sci. 8: 416-419. T h o m a s, R. L. 19 66. T he i n f l u e n ce of s e ed w e i g ht on s e e d l i ng v i g or in L o l i um p e r e n n e. Ann. Bot. 30: 111-121. T h r o n e b e r r y. G. O., and F. G. S m i t h. 19 55. R e l a t i on of t o ry and e n z y m a t ic a c t i v i ty to c o rn s e ed v i a b i l i t y. P h y s i o l. 30: 3 3 7 - 3 4 3. r e s p i r a- P l a nt T r u p p, C. R ., and I. T. C a r l s o n. 1971. I m p r o v e m e nt of s e e d l i ng v i g or of s m o o th b r o m e g r a ss ( B r o m us r e c u r r e nt s e l e c t i on f or high s e ed w e i g h t. C r o p. Sci. 11: 2 2 5 - 2 2 8. i n e r m is L e y s s .) by Vogel, W. C. 19 63. P l a n t i ng d e p th and s e ed s i ze e m e r g e n ce of b e a r d ed w h e a t g r a ss s e e d l i n g s. 16: 2 7 3 - 2 74 i n f l u e n ce J. R a n ge M g m t. W h a l l e y, R. D. B., and C. M. M c K e l l. 1967. I n t e r r e l a t i on of c a r- b o h y d r a te m e t a b o l i s m, s e e d l i ng d e v e l o p m e nt and s e e d l i ng g r o w th r a te of s e v e r al s p e c i es of P h a l a r i s. A g r o n. J. 59: 2 2 3 - 2 2 6. W h a l l e y, R. D. B., C. M. M c K e l l, and L. R. G r e e n. 1966a. S e e d- ling v i g or and t he n o n - p h o t o s y n t h e t ic s t a ge of s e e d l i ng g r o w th in g r a s s e s. C r o p. Sci. 6: 147-150. W h a l l e y, R. D. B., C. M. M c K e l l, and L. R. G r e e n. 1966b. E f f e ct of e n v i r o n m e n t al c o n d i t i o ns d u r i ng t he p a r e nt g e n e r a t i on on s e e d l i ng v i g or of t he s u b s e q u e nt s e e d l i n gs of O r y z o p s is m i l i a c ea ( L.) B e n t h. and Hook. C r o p. Sci. 6: 510-512. W h a l l e y, R. D. B., C. M. M c K e l l, and L. R. G r e e n. 1966c. S e ed p h y s i c al c h a r a c t e r i s t i cs and g e r m i n a t i on of h a r d i n g- g r a ss ( P h a l a r is ( H a c k .) H i t c h .) J. R a n ge MgmtT. 19: 129-132. t u b e r o sa v a r. s t e n o p t e ra W h y t e, R. O., T. R. G. M o i r, and J. P. C o o p e r. 1959. G r a s s es in A g r i c u l t u re F . A . O ., I t a l y. W i l l i a m s, R. F. 19 64. T he q u a n t i t a t i ve d e s c r i p t i on of g r o w t h. In G r a s s es and G r a s s l a n d s, C. B a n a rd (ed.) M a c m i l l a n, N ew Y o r k. pp. 89-101. W o o d s t o c k, L. W. 1969. B i o c h e m i c al t e s ts f or s e ed v i g o r. P r o c. Int. Seed. T e s t. A s s. 34: 2 5 3 - 2 6 3. W o o d s t o c k, L. W., and J. J. F e e l e y. 1965. E a r ly s e e d l i ng g r o w th and i n i t i al r e s p i r a t i on r a t es as p o t e n t i al i n d i c a t o rs of s e ed v i g or in c o r n. P r o c. A s s. Off. S e ed A n a l. 55: 131-139. 19 71. W r i g h t, L. N., and R. L. M a j o r. b l ue p a n i c g r a ss ( P a n i c um a n t i d ó t a le R e t z .) s e l e c t i o ns on shoot e l o n g a t i on and p r o d u c t i on of d ry m a t t e r. A g r o n. A b s t. 6 3 rd Ann. M e e t i ng A m e r. Soc. A g r o n. p. 17. I n f l u e n ce of s e ed weight of Z a v i t z, C. A. 1908. T he r e l a t i on b e t w e en t he s i ze of s e e ds and t he y i e ld of p l a n ts of f a rm c r o p s. P r o c. A m e r. Soc. A g r o n. 1: 98-105. D i s c u s s i on P e r i od M r. G a b e r t: U n d o u b t e d ly y o u ' ve o b s e r v ed m a ny s e e d l i ng v i g or t e s ts in t he l a b o r a t o r y. C an you m a ke a ny c o m m e n ts on t he d i- s e a s es or f u n gi t h at a re a p p a r e n t ly g r o w i ng on t he u n g e r m i n a t ed o r, in s o me c a s e s, t he g e r m i n a t ed s e e d s? A l s o, t he c o r r e s p o n d- ing e v e n ts w h i ch o c c ur i m m e d i a t e ly f o l l o w i ng or p r e c e e d i ng y o ur o b s e r v a t i o ns in t e r ms of w h e t h er t h e se f u n gi c a u s ed s e ed d e- t e r i o r a t i on or w h e t h er t he f u n gi c a me as an a f t er e f f e ct or s a p r o- p h y t e? D r. W h a l l e y: As you k n o w, t h e re a re a l a r ge n u m b er of m i c r o- f l o ra s p o r es and b a c t e r ia on t he o u t s i de of s e e d s. M a ny s e e ds a l so c a r ry a h e a vy e n d o g e n o us m i c r o f l o ra i n s i de t he e n d o s p e r m. I ' ve n o t i c ed w i th c r e s t ed w h e a t g r a ss and a l so with R u s s i an w i l d- r ye t h at t h e se e n d o g e n o us b a c t e r ia c an b e c o me v e ry i m p o r t a n t. If you m e a s u re t he r e s p i r a t i on r a te of one of t h e se s e e d s, you m ay f i nd a high r e s p i r a t i on r a t e. d i sh and let it g e r m i n a t e, t e s ta b r e a ks o p en and a m a ss of b a c t e r ia c o m es out and g r o w th s t o p s. S u r f a ce s t e r i l i z a t i on d o es not a f f e ct t h is e n d o g e n o us m a t e r i a l. I f e el in c e r t a in c i r c u m s t a n c es s o me s e ed l o ts c a r ry a v e ry h e a vy e n d o g e n o us m i c r o f l o ra a nd t h e se a re u n d o u b t e d ly i m p o r t a nt in t he f i e l d. it m ay g r ow f a st f or a d ay but t h en If you t h en put t he s e ed in a p e t ri t he T he cold t e st f or c o r n, w h i ch is u s ed as a v i g or t e s t, is c o n c e r n ed w i th r e s i s t a n ce to a t t a ck by m i c r o o r g a n i s m s. A ll t he t e st d o es is to s ee how r e s i s t a nt a s e ed lot is to s a p r o p h y t ic and p a t h o g e n ic m i c r o o r g a n i s ms p r e s e nt in t he s e ed or s o i l. U n d o u b t e d l y, I t h i nk t he m i c r o o r g a n i s ms a re m o st i m p o r t a n t, and p e r h a ps one could s e l e ct t he s p e c i es and s t r a i ns w i th a low e n d o g e n o us e n d o s p e rm m i c r o f l o r a. D r. Long: A f t er a c id s c a r i f i c a t i on and you go into a s t a n d a rd g e r m i n a t i on t e st in p e t ri d i s h, you n o t i ce a g r e at d e al m o re f u n g al a c t i v i t y. W e ' re w o n d e r i ng w h e t h er one a l t e rs t he e n d o s p e r m, or c h a n ge t he c a r b o h y d r a te p i c t u re h e re to w h e re c a r b on m a t e r i al is in a m o re a v a i l a b le f o r m; t h u s, e n c o u r a g i ng t he m i c r o o r g a n i s m s? is r e l a t ed to t he P r o b a b l y. S o me of t he r e s i s t a n ce to s a p r o p h y t ic D r. W h a l l e y: a nd f a c u l t a t i ve p a r a s i t ic m i c r o o r g a n i s ms l e a k a ge of c a r b o h y d r a t es f r om t he e n d o s p e rm of t he s e e ds d u r i ng g e r m i n a- t i o n. You c an s h ow t h is v e ry c l e a r ly w i th c o r n. m i n a t ed in o r d i n a ry p e t ri d i s h e s, t he w a t er f r om t he f i l t er pad c an be a n a l y z ed f or s u g a rs d u r i ng t he t e s t. S e ed l o ts w h i ch do w e ll in t he cold t e st u s u a l ly h a ve l ow s u g a rs in t he s o l u t i o n, low l e a k a ge f r om t he e n d o s p e rm and, t h e r e f o r e, l e ss e n e r gy s u b s t r a te f or t he m i c r o o r g a n i s m s. I would s u g g e st t h at t he a c id t r e a t m e nt in s o me c a s es i n c r e a s es t he l e a k i n e ss and, as a r e s u l t, t he g r o w th of f u n gi and b a c t e r i a. If s e e ds a re g e r- In a t u rf s e e d i ng e n v i r o n m e n t, D r. L o n g: is o ur m a j or p r o b l e m, what s i n g le t e st m i g ht be m o st i n d i c t i ve of g e t t i ng to s o m e t h i ng t h at would a p p l y? if a r a p id d r y i ng s e e d b ed If s e e d b ed d r y i ng is i m p o r t a n t, I would s c r e en D r. W h a l l e y: b r e e d i ng m a t e r i al f or t wo t h i n gs -- r a p i d i ty of g e r m i n a t i on and g e r m i n a t i on u n d er m o i s t u re s t r e s s. B o th of t h e se a re r e l a t i v e ly e a sy t e s ts to do. T he r a p i d i ty of g e r m i n a t i on is s i m p ly a m a t t er of c o u n t i ng t he n u m b er of s e e ds g e r m i n a t ed e a ch d ay in a s t a n d a rd g e r m i n a t i on t e s t. T he g e r m i n a t i on u n d er w a t er s t r e ss a l i t t le m o re e q u i p m e nt but is p o s s i b le on a r o u t i ne b a s i s. If you a r r a n ge an a p p r o p r i a te m e m b r a ne with a t h in l a y er of s o il a b o ve it ( l e ss t h an a c e n t i m e t er t h i c k) and a d i a l y s ed s o l u t i on of p o l y- e t h y l e ne g l y c ol ( 2 0 , 0 00 m o l e c u l ar w e i g h t) b e l o w, t he s o il m ay be k e pt at s ay - 5, - 10 and - 15 b a rs w a t er p o t e n t i al f or t he d u r a t i on of a g e r m i n a t i on t e s t. B r e e d i ng m a t e r i al c an t h en be s c r e e n ed f or p e r c e nt g e r m i n a t i on at d i f f e r e nt w a t er s t r e s s e s. r e q u i r es D r. D. T a y l o r: H a ve you a ny a d d i t i v es w h i ch m ay s p e ed g e r m i n a- t i on of s e e d s, or h a ve you r un a c r o ss a ny i n h i b i t o r s? T he t h at I a sk is t h at if y o u ' re p l a n t i ng a m i x t u re a nd one c o m p o n e nt s e e ms to d o m i n a t e, c an you s l ow down or s p e ed up t he g e r m i n a t i on a nd d e v e l o p m e nt of a ny one c o m p o n e n t? r e a s on I d o n 't k n ow of a ny g r o w th r e g u l a t o rs or c o m p o u n ds D r. W h a l l e y: t h at w i ll s p e ed up g e r m i n a t i on or s l ow it down in t he w ay t h at you a s k e d. T he only o n es t h at I am a w a re of a re t h o se w h i ch e i t h er b r e ak or i n d u ce s o me f o rm of s e ed d o r m a n c y. D r. F i s h e r: With r e s p e ct to y o ur l a ck of u n i f o r m i ty in y o ur s i m i l ar c o rn s e e d s, one of o ur p l a nt b r e e d e rs b r o u g ht me s o me oat s e ed t h at he had s e p a r a t ed into t he p r i m a ry s e ed in t he s p i k e l et and t he s e c o n d a r y, w h i ch w e re s c r e e n ed f or i d e n t i c al w e i g h t s. Yet, t he p r i m a ry g e r m i n a t ed and g r ew m o re r a p i d ly t h an t he s e c o n- d a r y. W h en t h e se w e re s e c t i o n e d, we f o u nd t h at i n v a r i a b ly t he p r i m a ry s e ed h ad one m o re f o l i ar r i d ge in t he e m b r yo t h an t he s e c o n d a ry one. T h is s h o ws up v e ry m a r k e d ly in o a t s, w h i ch is a p a n i c le g r a s s. I s u s p e ct in b l u e g r a ss it m u st be v e ry i m p o r t a n t. D r. W h a l l e y: Yes, I e n t i r e ly a g r ee with you. T h e re h as b e en s o me w o rk w h e re p e o p le h a ve e i t h er by m a n i p u l a t i on or s e l e c t i on o b t a i n ed s e e ds of s i m i l i ar t o t al s i ze w e i g h t, but b i g g er and s m a l l er e m b r y o s, and f o u nd e x a c t ly t he s a me e f f e ct t h at you m e n t i o n e d. T h o se w i th l a r g er e m b r y os s e em to get a b e t t er s t a rt and g r ow f a s t e r. I d o n 't k n ow h ow long t he a d v a n t a ge c o n t i n u e s, or h ow good t he c o r r e l a t i on is b e t w e en e m b r yo s i ze or leaf n u m b er and s e e d l i ng s u r v i v al or any p l a nt s i ze c h a r a c t e r i s t ic at s ay s ix or eight w e e ks a f t er g e r m i n a t i o n. w h e a t g r a s s, but t h e re m ay be b e t t er c o r r e l a t i on with o t h er I t h i nk t h e re is a p o or c o r r e l a t i on in c r e s t ed s p e c i e s. D r. Yu: F or d i f f e r e nt p l a nt s p e c i e s, how would you d e f i ne t he s e e d l i ng s t a g e? D r. W h a l l e y: T he d e v e l o p m e nt of a p l a nt f r om t he t i me of s e ed i m b i b i t i on to t he t i me of s e ed p r o d u c t i on is e s s e n t i a l ly a c o n t i n u u m. F or c o n v e n i e n ce one t e n ds to d i v i de b er of s t a g e s, but I t h i nk it is i m p o r t a nt to r e m e m b er t h at s u b d i v i s i o ns a re e s s e n t i a l ly a r b i t r a r y. T h e r e f o r e, t he end of t he s e e d l i ng s t a ge is w h en you d e f i ne it. B e c a u se of my i n t e r e st s e e d l i ng s u r v i v a l, I ' w o u ld p e r h a ps u se a d e f i n i t i on b a s ed on m o r- t a l i ty t h is c o n t i n u um into a n u m- t h e se r a t e s. in D r. F i s h e r: I would s ay t h at t he s e e d l i ng s t a ge is o v er w h en t he a p i c al m e r i s t em m a k es i ts f i r st n ew f o l i er r i d g e. But t h a t 's a r- b i t r a r y. D r. W h a l l e y: Y e s. D r. Long: c o m m e nt on h is o b s e r v a t i o ns m i n a t i on r a t es in r e l a t i on to n u t r i e n t s. in t e r ms of I t h i nk it would be i n t e r e s t i ng to h a ve D r. N i t t l er v a r i e t i es as to g e r- I h a v e n 't r e a l ly s t u d i ed t h i s. D r. N i t t l e r: I do k n ow t h at s o me v a r i e t i es h a ve m u ch l a r g er s e e ds t h an o t h e r s. i n t e r e s t ed if s o m e b o dy would r un a c o r r e l a t i on b e t w e en t he s e e d l i ng e s t a b- l i s h m e nt and v a r i e t i es t h at h a ve d i f f e r e nt s e ed s i z e. I 'd be D r. Long: Did you n o t i ce a ny d i f f e r e n ce in y o ur w o rk w h en you h ad m i n us c a l c i um v e r s us c a l c i u m? Did s o me g e r m i n a t e? A l s o, did t he r a te of s e e d l i ng g r o w th r a te v a r y? D r. N i t t l e r: No, I h a v e n 't r e a l ly n o t i c ed a ny d i f f e r e n c e, we get p o or g e r m i n a t i on w i th s o me s o l u t i o n s. It v a r i es with s e ed l o ts s o m e t i m e s. We t e s t ed s e v e r al s a m p l es of W i n d s or and f o u nd t h at s o me s a m p l es t h at g e r m i n a t ed p o o r ly o t h e r w i se w i ll g e r m i n a te r e l a t i v e ly w e ll if p o t a s s i um n i t r a te is u s e d. But we h a v e n 't c o r- r u l a t ed t h is with s e e d l i ng g r o w t h. t h o u gh D r. P a y n e: You h ad a c a l c i u m - n i t r o g en i m b a l a n ce t h e re t h at w as c a u s i ng a d i f f e r e n t i a l? D r. N i t t l e r: Y e s, I t h i nk t h is is p a rt of it. I D r. M a d i s o n: T h is is not a q u e s t i o n, but a c o m m e n t a r y. h a ve l o o k ed at P e n n c r o ss f r om t he s t a n d p o i nt of c o m p e t i t i on a f f e c t i ng e s t a b l i s h m e n t. A f t er 14 p h y t o c h r o n s, my w e a k er p l a n ts u n d er t he s t r o ng s t r e ss of c o m p e t i t i on had m a de 14 p h y t o m e r s, of w h i ch 4 w e re s t i ll f u n c t i o n i n g. At t he o t h er e x t r e m e, with no c o m p e t i t i o n, I ' ve had v i g o r o us p l a n ts with o v er 1,000 p h y t o m e rs a f t er 14 p h y t o c h r o n s, and w i th as m a ny as 1,400 p h y t o m e r s. In t h e se e x t r e m e ly v i g o r o us p l a n t s, t he p r o p h y ll h as a bud w h i ch p r o d u c es a s e c o nd p r o p h y ll and s e c o nd b r a n ch w i t h in t he f i r st p r o p h y l l. T h is o b s e r v a t i on s h o ws a c h a r a c t e r i s t ic of t he e x t r e m e ly v i g o r o us p l a n t; t he p r o p h y ll h as a bud. P E N N F I NE P E R E N N I AL R Y E G R A SS D E S C R I P T I ON AND P E R F O R M A N CE C H A R A C T E R I S T I CS J. M. D u i ch and G. W. M c K ee D e p a r t m e nt of Agronomy- P e n n s y l v a n ia S t a te U n i v e r s i ty U n i v e r s i ty P a r k, P e n n s y l v a n ia 16802 I n t r o d u c t i on U n d er t he p r o v i s i o ns of t he 19 70 P l a nt V a r i e ty P r o t e c t i on Act, t he p r o b l em of p r o v i d i ng an o b j e c t i ve d e s c r i p t i on of a n o v el v a r- i e ty h as b e c o me m o st c r i t i c a l. T he a b s e n ce of r e f i n ed b i o c h e m- i c al and a s s o c i a t ed t e c h n i q u es s t i ll n e c e s s i t a t es r e l i a n ce on t r a- d i t i o n al m o r p h o l o g i c al and u se p e r f o r m a n ce d a t a. T h is p a p er d e a ls w i th t he m e t h o ds u s ed to d e s c r i be P e n n- f i ne p e r e n n i al r y e g r a ss ( L o l i um p e r e n ne L.) a nd is i n t e n d ed as an aid to o t h e rs f a c i ng t he p r o b l em of p r o v i d i ng v a r i e t al d e s c r i p t i o n s. O r i g in and B r e e d i ng H i s t o ry P e n n f i ne r y e g r a ss is a n o n f l u o r e s c e nt t u r f - t y pe p e r e n n i al r y e g r a ss d e v e l o p ed as a t h r e e - c l o ne s y n t h e t ic v a r i e ty at T he P e n n- s y l v a n ia A g r i c u l t u r al E x p e r i m e nt S t a t i on ( D u i c h, P e r k i n s, and Cole, 19 72). A ll t h r ee c l o n es w e re s e l e c t ed in S o u t h e a st P e n n- s y l v a n i a; t wo a re golf c o u r se f a i r w ay s e l e c t i o n s, and t he t h i rd w as f o u nd on a t e n n is c o u r t. O r i g i n al s e ed s o u r c es a re u n k n o wn and t he a r e as w e re s e e d ed and r e s e e d ed as t u rf f or a p p r o x i m a t e ly 40 y e a r s. A ll c l o n es in t he r y e g r a ss p r o g r am r e q u i r ed a s p r e ad of 12 i n c h es m i n i m u m, and the c l o n es in q u e s t i on r a n g ed f r om 16 to 28 i n c h e s. T h is f a ct l e a ds one to e s t i m a te an a ge of at l e a st 15 y e a rs u n d er c l o s e - c ut turf s e l e c t i on p r e s s u r e. B r e e d e r 's s e l e c t i on i n v o l v ed t e x t u r e, m o w a b i l i t y, w i n t er v i v a l, s n o w m o ld t o l e r a n c e, and h e a d i ng and a n t h e s is d a t e s. to t u rf q u a l i ty t e s t i n g, u n i f o r m i ty of h e a d i ng and a n t h e s is w e re a l so c o n f i r m ed in two y e a rs of t e s t i ng in O r e g on u n d er c o m m e r- c i al p r o d u c t i on c o n d i t i o n s. s u r- P r i or T he P e n n f i ne v a r i e ty s h o ws a h i gh d e g r ee of u n i f o r m i ty f or a t h r e e - c l o ne s y n t h e t i c. U n i f o r m i ty t e s t i ng i n v o l v ed two g e n e r a- t i o ns of s p a ce p l a n t ed and t u rf t r i a l s. T wo F o u n d a t i on g r o w e rs f a i l ed to d e t e ct d i s t i n c t i ve v a r i a n ts in 40 a c r es of p r o d u c t i on in O r e g on and W a s h i n g t o n. O b s e r v a t i o ns to d a te i n d i c a te P e n n f i ne to be a r e l a t i v e ly s t a b le v a r i e ty on t he b a s is of m o r p h o l o g i c al c h a r a c t e rs and t u rf r e s p o n s e. C o m m e r c i al p r o d u c t i on w i ll i n v o l ve t h r ee g e n e r a t i o n s: B r e e d e r s, F o u n d a t i on and C e r t i f i e d. B r e e d e r s' s e ed will be p r o d u c ed at U n i v e r s i ty P a r k, P a ., and o t h e rs in t he P a c i f ic N o r t h w e st s t a t e s. C o m m e r c i al s e ed w i ll be in t he " c e r t i f i ed o n l y" c a t e g o r y, w i th s e ed f a i l i ng to m e et c e r t i f i ed s t a n d a r ds not e l i g i b le f or v a r i e ty n a m e. M a t u re P l a nt C o m p a r i s o ns To c o m p a re p l a n ts of p e r e n n i al r y e g r a s s, m e a s u r e m e n ts w e re m a de in l a te J u ne 19 71 of f i f ty p l a n ts e a ch of s ix c u l t i v a rs of p e r e n n i al r y e g r a ss g r o w i ng in W e s t e rn Idaho on t he R a t h- d r um R a n ch of t he J a c k l in S e ed C o m p a n y. T he r a n ch is l o c a t ed a p p r o x i m a t e ly 20 m i l es e a st of S p o k a n e, W a s h i n g t o n. E a ch plot of r y e g r a ss w as a p p r o x i m a t e ly 5 m long by 2 m wide. P l o ts h ad b e en f e r t i l i z ed and i r r i g a t ed as w as c u s t o m a ry f or s e ed p r o d u c t i on in t h is a r e a. A ll p l a n ts m e a s u r ed had h e a d ed and a n t h e s is had o c c u r r e d. S e ed would p r o b a b ly h a ve b e en r i pe 10 to 14 d a y s. P l o ts had b e en e s t a b l i s h ed in m i d - S e p t e m b er 19 70 to e v a l u a te r y e g r a ss c u l t i v a rs u n d er e n v i r o n m e n t al c o n d i- t i o ns f or c o m m e r c i al s e ed p r o d u c t i o n. in M e a s u r e m e n ts w e re m a de of f l a g l e af s i z e, h e ad length, i n- t e r n o de l e n g th and h e ad weight. D a ta w e re a n a l y z ed by a n a l y s is of v a r i a n ce and s i g n i f i c a nt d i f f e r e n c es d e l i n e a t ed by D u n c a n 's m o d i f i ed ( B a y e s i a n) l e a st s i g n i f i c a nt d i f f e r e n ce t e s t. H e ad w e i g ht To c o m p a re h e ad w e i g h t, f i ve r e p l i c a t i o ns of 15 h e a ds e a ch of s ix c u l t i v a rs of p e r e n n i al r y e g r a ss w e re w e i g h ed to 0.01 g. T he h e a ds w e re a i r - d r i ed and t r i m m e d, w i th s t e ms b e l ow t he l o w e st f l o r et r e m o v e d. H e a ds of P e n n f i ne w e i g h ed t he l e a st and t h o se of N o r l e a, t he m o st ( T a b le 1). T he c o e f f i c i e nt of v a r i a b i l i ty f or t h is m e a s u r e m e nt w as 6.2%. H e ad l e n g th To c o m p a re h e ad length, 50 h e a ds of e a ch of s ix c u l t i v a rs of p e r e n n i al r y e g r a ss w e re m e a s u r ed to t he n e a r e st m i l l i m e t e r. T he h e a ds h ad b e en a i r - d r i e d. M e a s u r e m e n ts w e re m a de f r om t he t ip of t he h e ad to t he of P e n n f i ne w e re t he s h o r t e st a nd t h o se of N o r l e a, t he l o n g e st ( T a b le 2). T he c o e f f i c i e nt of v a r i a b i l i ty f or t h is m e a s u r e m e nt w as 14.8%. i n t e r n o de b e l ow t he l o w e st f l o r e t. H e a ds F l a g l e af To c o m p a re s i ze of t he f l a g l e af in s ix c u l t i v a rs of p e r e n n i al r y e g r a s s, m e a s u r e m e n ts of t he f l a g l e af l e n g th and w i d th w e re m a de on 50 p l a n ts e a ch of s ix c u l t i v a rs of p e r e n n i al r y e g r a s s. F l a g l e af w i d th to t he n e a r e st 0.1 mm w as m e a s u r ed at a spot 4 mm f r om t he b a s al end w h e re t he leaf j o i n ed t he leaf s h e a t h. M e a s u r e- m e n ts w e re m a de with a 10X m a g n i f y i ng g l a ss e q u i p p ed w i th a r e t i c le c a l i b r a t ed to 0.1 m m. F l a g l e af l e n g th w as m e a s u r ed to t he n e a r e st 0.5 mm u s i ng a s t e el m e t r ic t a p e. L e n g th m e a s u r e m e n ts w e re m a de f r om leaf t ip to a point w h e re t he leaf and leaf s h e a th j o i n e d; i . e ., on a l i ne with t he l i g u l e. T he f l a g l e af of P e n n f i ne w as by f ar t he s h o r t e st and n a r r o w e s t, w h i le t h at of N o r l ea w as t he l o n g e st and w i d e st ( T a b le 3). T he d e r i v ed m e a s u r e m e nt LW a c c e n t u a t ed leaf m e a s u r e m e nt d i f f e r e n c es so t h at a ll s ix c u l t i v a rs w e re s i g n i f i c a n t ly d i f f e r e n t, one f r om t he o t h e r. f e r e n c es in t he r a t io of f l a g l e af l e n g th to w i d th ( L / W) w e re not as d i a g n o s t ic of c u l t i v ar d i f f e r e n c es as m e a s u r e m e n ts of f l a g l e af l e n g t h, w i d th a nd LW. T he c o e f f i c i e n ts of v a r i a b i l i ty f or m e a s u r e- m e n ts of f l a g l e af l e n g th and w i d th w e re 17.6 and 14.2%, F or t he d e r i v ed q u a n t i t i es LW and L / W, t he c o e f f i c i e n ts of v a r i a- b i l i ty w e re 25.9 and 19.2%, In c o n t r a s t, d i f- r e s p e c t i v e l y. r e s p e c t i v e l y. I n t e r n o de l e n g th I n t e r n o de l e n g th w as m e a s u r ed to t he n e a r e st To c o m p a re i n t e r n o de l e n g t h, m e a s u r e m e n ts to t he n e a r e st m i l l i m e t er w e re m a de of t he f i r st ( b a s a l) s ix i n t e r n o d es of t he r y e g r a ss h e a d. millimeter. For e a ch internode, P e n n f i ne w as t he s h o r t e s t. I n- t e r n o d es of Linn w e re e i t h er t he l o n g e st ( i n t e r n o d es 2, 3) or not s i g n i f i c a n t ly d i f f e r e nt f r om t he l o n g e st ( i n t e r n o d es 4, 5). T he c o e f f i c i e n ts of v a r i a b i l i ty f or i n t e r n o de l e n g th r a n g ed f r om 21.3 to 2 5 . 7 %. T he r a t i os of one i n t e r n o de to a n o t h er w e re a l so c o m p a r ed ( T a b le 5). S o me v a r i e t i es d i f f e r ed s i g n i f i c a n t ly (p = .05) f r om o t h e rs in t he r a t i os of e a ch i n t e r n o de l e n g th to i n t e r n o d es 1 and 2. R a t i os of t he l e n g th of e a ch i n t e r n o de to i n t e r n o d es 3, 4, 5 and 6 w e re g e n e r a l ly n o n s i g n i f i c a n t. T he c o e f f i c i e n ts of v a r i a b i l i ty f or t he l e n g th of e a ch i n t e r n o de to i n t e r n o de 1 v a r i ed f r om 17.3 to 28.6%; f or t he r a t i os to i n t e r n o de 2, f r om 4 6 .6 to 68.0%, and f or t he r a t i os to i n t e r n o de 3, f r om 67.8 to 102.3%. R a n k i ng of s ix c u l t i v a rs f r om l e a st to g r e a t e st f or p l a nt t he r e l a t i ve r a n k i ng w a s: P e n n f i n e, 12; M a n- c h a r a c t e r i s t i c s, h a t t a n, 29; Linn, 43; P e l o, 52; N o r l e a, 55, and NK-100, 58 ( T a b le 6). T h u s, as t he a b o ve d a ta i n d i c a t e, P e n n f i ne h as s m a l l er s h o r t er and l i g h t er h e a d s, a nd s h o r t er f i ve c u l t i v a rs e v a l u a t e d. i n t e r n o d es t h an t he o t h er f l a g l e a v e s, R e l a t i ve m a t u r i ty O b s e r v a t i o ns J u ne 3 and 16, 19 71, i n d i c a t ed t h at w i th r e g a rd to r e l a t i ve m a t u r i ty a nd h e ad e m e r g e n c e, P e n n f i ne and L i nn would be c l a s s i f i ed as e a r l y, N K - 1 00 as m e d i u m, N o r l ea as m e d i um l a t e, a nd P e lo and M a n h a t t an as l a te ( T a b le 7). S e ed c h a r a c t e r i s t i cs No d i s c e r n i b le c h a r a c t e r i s t i cs of P e n n f i n e. r y e g r a ss s e ed h a ve b e en d e t e c t ed to d a t e, but w o rk is c o n t i n u i ng along t h e se l i n e s. S e ed s i z e, n u m b er and weight a p p e ar to be a p p r e c i a b ly s m a l l er and l i g h t er t h an Linn, and s i m i l ar to M a n h a t t a n. T h e se d a ta c a n n ot be f i n a l ly a s s e s s ed u n t il a r e p r e s e n t a t i ve n u m b er of s a m p l es a re o b t a i n ed f r om p r o d u c t i on a r e as to d e t e r m i ne a nd y e ar v a r i a b l e s. l o c a t i on T r i al P e r f o r m a n ce P e n n f i ne r y e g r a ss w as d e v e l o p ed f or t u r f g r a ss u s e. T u r f- i n i t i a t ed in 19 65 at U n i v e r s i ty P a r k, P a ., c o m p a r ed g r a ss t r i a ls v a r i o us c u l t i v a rs u n d er a c c e p t a b le m a i n t e n a n ce p r o c e d u r e s. S e e d i n gs w e re m a de at 4 p o u n ds p er t h o u s a nd a nd m a i n t a i n ed at 1 l / 4 - i n ch cutting h e i g h t. F e r t i l i ty w as at t he m o d e r a te l e v el of 3 p o u n ds of n i t r o g en p er t h o u s a n d. s t r e ss t r e a t m e nt with a l d r in and a n n u al a p p l i c a t i o ns of p h e n o xy c o m p o u n ds f or b r o a d l e af w e ed c o n t r o l. l e v e ls only. C h e m i c al a p p l i c a t i o ns w e re l i m i t ed to g r ub I r r i g a t i on w as a p p l i ed at wilt T u r f g r a ss p e r f o r m a n ce d a ta in U n i v e r s i ty P a rk t r i a ls a re s h o wn in T a b l es 1 to 4. A ll e v a l u a t i o ns f or d e n s i t y, m o w i ng q u a l i ty a nd d i s e a se w e re m a de by t wo to f o ur o b s e r v e r s. S e e d l i ng g r o w th w as m e a s u r ed and c o m p a r ed to t he q u i ck g r o w i ng a n n u al as s t a n d a rd in f o ur m o w i n gs f o l l o w i ng e m e r g e n c e. Leaf w i d th m e a s u r e- m e n ts w e re m a de on 50 f i r st and s e c o nd l e a v es b e l ow budleaf p er plot with a m i c r o m e t e r. F l u o r e s c e n ce d e t e r m i n a t i o ns w e re m a de at T he P e n n s y l v a n ia D e p a r t m e nt of A g r i c u l t u re S e ed L a b o r a t o r y. t he r D e n s i ty P e n n f i ne and M a n h a t t an r y e g r a s s es p e r s i s t ed at t he h i g h e st d e n s i ty l e v e ls a m o ng n i ne c u l t i v a rs o v er a 5 - y e ar p e r i od ( T a b le 8). U t i l i z i ng a 1 to 10 r a t i ng s c a le w i th 10 : b e s t, P e n n f i ne and M a n- h a t t an e x c e e d ed 8.5 f or s ix e v a l u a t i ng p e r i o d s, and o u t - p e r f o r m ed a ll o t h er e n t r i e s. Mowing q u a l i ty A d i s t i n g u i s h i ng f e a t u re of P e n n f i ne is t he m o w a b i l i ty or r e- l a t i ve a b s e n ce of f o l i ar s h r e d d i ng f o l l o w i ng m o w i ng with a r e el m o w er ( T a b le 9). T h is is a w e l l - r e c o g n i z ed p r o b l em of d u r i ng high t e m p e r a t u re p e r i o d s. With one e x c e p t i o n, P e n n f i ne m a i n t a i n ed a m o w i ng q u a l i ty a b o ve 8.0 f or 5 y e a r s, a v e r a g i ng 8.3. O t h er c u l t i v a rs r a n g ed f r om 2.6 to 7.3. r y e g r a s s es D i s e a se t o l e r a n ce N a t u r al d i s e a se i n f e c t i o ns of e p i d e m ic m a g n i t u de i n c l u d ed ( f o l i a r ), leaf r u st and H e l m i n t h o s- s n o w m o l d, F u s a r i um r o s e um p o r i um ( s p e c i es u n i d e n t i f i a b le by c u l t u r e) ( T a b le 10). Of n i ne c u l t i v a r s, P e n n f i ne r e s i s t a n ce r a n k ed f i r st f or s n o w m o l d, t h i rd f or F u s a r i u m, f o u r th f or r u s t, and f i r st f or H e l m i n t h o s p o r i u m. A d d i t i o n al s n o w m o ld and w i n t er s u r v i v al d a ta w e re o b t a i n ed f r om f o r a ge plot s e e d i n gs at t he U.S. P a s t u re L a b o r a t o ry ( T a b le 11). F l u o r e s c e n c e, v e r t i c al g r o w th and t e x t u re A ll c u l t i v a rs w e re a n a l y z ed f or p u r i t y, g e r m i n a t i on and f l u o r e s c e n ce by T he P e n n s y l v a n ia S t a te S e ed L a b o r a t o ry ( T a b le 12). P e n n f i n e, M a n h a t t a n, P e l o, N o r l ea and C o m bi s h o w ed z e ro f l u o r- e s c e n c e, w h e r e as o t h er p e r e n n i a ls r a n g ed f r om 3.5 to 5.0%. F o ur m e a s u r e m e n ts f o l l o w i ng s e e d i ng in 19 65 s h o w ed P e n n- f i ne h ad t he l e a st v e r t i c al g r o w t h, a v e r a g i ng 40% of a n n u al g r a s s. M a n h a t t an a v e r a g ed 50%, and o t h er c u l t i v a rs r a n g ed f r om 60 to 80%. Low v e r t i c al g r o w th is v e ry d e s i r a b l e, p a r t i c u l a r ly r y e g r a s s es in t e r ms of c o m p e t i t i on and m o w i ng f r e q u e n cy w h en a re u s ed as c o m p a n i on g r a s s es with s l o w er d e v e l o p i ng s p e c i e s, s u ch as b l u e g r a s s. r y e- L e a f - w i d th m e a s u r e m e n ts d u r i ng t he s e c o nd g r o w i ng s e a s on ( 19 67) s h o w ed P e n n f i ne h ad t he f i n e st t e x t u re of t he n i ne c u l t i v a r s. P e n n f i ne a v e r a g ed 2.5 mm f or t he f i r st m a t u re c u l t i v a rs r a n g ed f r om 2.7 to 5.6. T he v a r i e ty n a me w as s u g g e s t ed by t h is t e x t u r al c h a r a c t e r. l e a f, and o t h er O v er s e e d i ng V a r i e t al d i s t i n c t i on is s u p p o r t ed by b e r m u d a g r a ss o v e r s e e d i ng t r i a ls at S t a te C o l l e g e, M i s s ., and C o l l e ge Station, T e x ., as s h o wn in T a b l es 13 and 14, r e s p e c t i v e l y. B o th s t a t i o ns r e p o rt a u n i q ue h i gh t e m p e r a t u re p e r s i s t a n ce d u r i ng t he l a t t er p a rt of t he s e a s o n. P e n n f i ne h as a l so p e r s i s t ed u n d er high t e m p e r a t u re c o n d i t i o ns in S o u t h e rn C a l i f o r n i a, as n o t ed in a p e r s o n al l e t t er f r om Stan S p a u l d- ing, U n i v e r s i ty of C a l i f o r n i a, South C o a st E x p e r i m e nt Station, S a n ta Ana. R e f e r e n c es C i t ed D u i c h, J. M., A. T. P e r k i n s, H. C o l e. 1972 R e g i s t r a t i on of P e n n- f i ne P e r e n n i al R y e g r a s s. C r op Sci. 12:257. T a b le 1. C o m p a r a t i ve w e i g ht of 15 h e a ds of s ix c u l t i v a rs of p e r- e n n i al r y e g r a s s. V a r i e ty P e n n f i ne M a n h a t t an N K - 1 00 L i nn P e lo N o r l ea Weight, g 1.50 a* 2.05 b 2. 19 be 2 . 23 be 2.56 c 3.27 d * M e a ns f o l l o w ed by t he s a me l e t t er do not d i f f er s i g n i f i c a n t ly at p - .01 by D u n c a n 's m o d i f i ed ( B a y e s i a n) l e a st s i g n i f i c a nt d i f- f e r e n ce t e s t. T a b le 2. C o m p a r a t i ve h e ad l e n g th of s ix c u l t i v a rs of p e r e n n i al r y e g r a s s. V a r i e ty P e n n f i ne Linn M a n h a t t an P e lo N K - 1 00 N o r l ea H e ad l e n g t h, cm 16.73 a* 19.72 b 2 3 . 44 c 25.05 d 2 5 . 93 d 30.22 e ' M e a ns f o l l o w ed by t he s a me l e t t er do not d i f f er s i g n i f i c a n t ly at p = .01 by D u n c a n 's ( B a y e s i a n) l e a st s i g n i f i c a nt d i f f e r e n ce t e s t. T a b le 3. C o m p a r a t i ve s i ze of f l a g l e af of s ix c u l t i v a rs of p e r e n n i al r y e g r a ss ( p l a n ts h e a d e d ). L e n g th ( L), Width (W), V a r i e ty mm P e n n f i ne L i nn M a n h a t t an P e lo N K- 100 N o r l ea 138.6 a* 175.9 b 206.7 c 255.8 d 262.5 d 282.2 e mm 4.29 a 5.38 b 5.29 b 6.26 c 5.52 b 6.55 c LW, mm L /W 603.58 a 9 4 9 . 87 b 1102.12 c 1610.99 e 1460.18 d 1858.21 e 32.66 a 33.47 a 39.60 b 4 1 . 11 be 4 7 . 96 d 43.72 c ^ R e a d i ng v e r t i c a l ly f or e a ch c o m p a r i s o n, m e a ns f o l l o w ed by t he s a me l e t t er do not d i f f er s i g n i f i c a n t ly at P = .01 by D u n c a n 's m o d i f i ed ( B a y e s i a n) l e a st s i g n i f i c a nt d i f f e r e n ce t e s t. T a b le 4. C o m p a r a t i ve i n t e r n o de l e n g th of h e a ds of s ix c u l t i v a rs or p e r e n n i al r y e g r a s s. I n t e r n o de n u m b er 1 2 3 mm 4 5 1 24.0 a* 25.0 a 28.2 b 31.8 c 32.4 c 34.5 c 20.2 a 21.6 a 25.2 b 2 7 .5 b 26.8 b 2 7 .7 b 15.6 a 17. 1 a 17.5 a 2 0 .3 b 21.3 b 21.6 b 13.6 a 15.4 b 16.9 be 18.6 cd 20.0 d 17.9 c 12.1 a 14.1 b 14.0 b 14.8 be 17. 5 d 16.1 cd 11.1 a 12.4 ab 13.6 b 13.2 b 15.6 c 13.2 b V a r i e ty P e n n f i ne M a n h a t t an N o r l ea P e lo N K - 1 00 L i nn * M e a ns f o l l o w ed by t he s a me l e t t er do not d i f f er s i g n i f i c a n t ly at p = .01 by D u n c a n 's m o d i f i ed ( B a y e s i a n) l e a st s i g n i f i c a nt d i f- f e r e n ce t e s t. T a b le 5. C o m p a r a t i ve i n t e r n o de r a t i o s, h e a ds of s ix c u l t i v a rs of p e r e n n i al r y e g r a s s. I n t e r n o de r a t io V a r i e ty 2 /l 3 /l 4 /l 5 /l 6 /l L i nn N K - 1 00 P e n n f i ne M a n h a t t an P e lo N o r l ea .82 a* .84 ab .85 ab .87 ab .88 ab .90 b .64 a .67 a .60 a .69 a .60 a .63 a .54 a .63 b .58 ab .62 b .60 b .62 b .49 a .56 bc .52 a bc .57 c .49 a .50 ab .39 a .50 c .47 bc .50 c .43 ab .50 c I n t e r n o de r a t io 1 /2 3 /2 4 /2 5 /2 6 /2 L i nn N K - 1 00 P e n n f i ne M a n h a t t an P e lo N o r l ea 1.25 a 1.45 a 1.20 a 1. 17 a 1.18 a 1.18 a .79 a .96 a .78 a .80 a .76 a . 73 a .66 a .85 b .68 a .72 ab .70 ab .69 a .59 ab .79 b .61 ab .66 ab .56 a .60 ab .48 a .68 b .56 ab .59 ab .50 a .55 ab * F or e a ch r a t io c o m p a r i s o n, c u l t i v ar m e a ns f o l l o w ed by t he s a me l e t t er do not d i f f er at p z .05 by D u n c a n 's m o d i f i ed ( B a y e s i a n) l e a st s i g n i f i c a nt d i f f e r e n ce t e s t. T a b le 6. C o m p a r a t i ve r a nk o r d er f or 12 c h a r a c t e r i s t i cs of s ix c u l t i v a rs of p e r e n n i al r y e g r a s s. H e ad F l a g l e af I n t e r n o de l e n g th V a r i e ty W i d th L e n g th L e n g th Width LW L /W 1 2 3 4 5 6 P e n n f i ne M a n h a t t an L i nn P e lo N o r l ea N K - 1 00 1* 2 4 5 6 3 1 3 2 4 6 5 1 3 2 4 6 5 1 2 3 5 6 4 1 3 2 5 6 4 1 3 2 4 6 5 1 1 1 1 1 1 2 2 2 2 3 2 6 6 6 4 5 4 4 5 4 5 4 3 3 3 3 3 2 5 5 4 5 6 6 6 ^ R a n k ed in a s c e n d i ng o r d er f r om l e a st ( 1) to g r e a t e st (6). T a b le 7. C o m p a r a t i ve t i me to h e a d, s ix c u l t i v a rs of p e r e n n i al r y e g r a s s .* R e p r o d u c t i on s t a t us V a r i e ty J u ne 3 J u ne 16 h e a d ed h e a d ed s t a r t i ng to h e ad L i nn P e n n f i ne N K - 1 00 N o r l ea P e lo M a n h a t t an h e a d ed s t a r t i ng to h e ad s t a r t i ng to h e ad R e l a t i ve m a t u r i ty e a r ly e a r ly m e d i um m e d i um l a te l a te l a te * B a s ed on o b s e r v a t i o ns by J a c k l in S e ed C o m p a ny p e r s o n n e l. T a b le 8. R y e g r a ss t u rf e v a l u a t i on -- d e n s i t y, U n i v e r s i ty P a r k, P a. V a r i e ty 7 / 1 5 / 66 9 / 1 6 / 66 7 / 2 7 / 67 8 / 2 9 / 67 9 / 4 / 68 8 / 9 / 69 D e n s i ty r a t i n g* P e n n f i ne M a n h a t t an N K- 100 P e lo N o r l ea B r a n b a n t ia C o m bi Linn A n n u al LSD .05 8.8 8.7 8.0 8.2 7.7 8.3 9 .0 6.7 1.7 1.1 9 .0 8.6 7.0 7.3 5.8 7.3 8.0 6.8 1.0 1.3 * S c a le 1 to 10; 10 : b e s t. 9 .1 9 .1 8.3 8.7 7.7 8.0 8.0 7.7 1.2 1.6 8.9 9 .0 7.8 8.7 6.8 8.2 8.3 7.0 — 1.4 9 .0 9 .0 7.0 7.3 6.2 7.2 7.3 5.7 — 1.7 8.8 8.7 6.5 7.2 6.3 6.0 6.2 5.3 — 1.9 T a b le 9. R y e g r a ss t u rf e v a l u a t i on -- m o w i ng q u a l i t y, U n i v e r s i ty P a r k, P a. V a r i e ty 7 / 1 5 / 66 8 / l l / 66 8 / 2 7 / 67 7 / 2 8 / 68 8 / 9 / 69 A v e r a ge Mowing q u a l i t y* P ennf i ne M a n h a t t an NK - 100 P e lo N o r l ea B r a n b a n t ia C o m bi L i nn A n n u al LSD .05 9 .0 8.0 4.0 5.3 8.0 6.0 6.0 4.7 2.0 1.2 8.0 7.0 3.0 3.3 5.7 4 .3 3.3 2 .3 — 1.9 8.2 7.5 4 .5 4 .5 5.5 5.5 5.0 2.0 — 1.3 * S c a le 1 to 10; 10 : b e s t. 8.3 7.7 5.7 6.3 7.0 6.7 6.3 2.0 — 1.4 7.8 6.2 4.7 6.7 6.7 4 .7 4 .7 2.2 — 1.3 8.3 7.3 4 .4 5.2 6.6 5.4 5.1 2.6 — T a b le 10. R y e g r a ss t u rf e v a l u a t i on -- d i s e a s e, U n i v e r s i ty P a r k, P a. % s n o w m o ld F. % s n o w m o ld r o s e u m* r u s t* H e l m i n t ho s p o r i um r u s t* s p p .* V a r i e ty V a r i e ty V a r i e ty 1966 1970 1966 1967 1968 19 69 P e n n f i ne M a n h a t t an N K - 1 00 P e lo N o r l ea 6 8 7 8 10 B r a n b a n t ia 12 C o m bi L i nn A n n u al 10 10 55 2 8 37 13 20 - 30 43 99 LSD .05 13 17 -¡'Scale 1 to 10; 1 - l e a s t. 3.3 2.0 6.0 5.0 6 .3 6.0 5.3 5.0 2.0 1.4 2.0 1.2 2.0 2.0 7.0 1.5 3.0 1.0 1.7 6.0 5.0 5.0 4.0 3.7 4 .3 2 .3 2.1 2.0 2.1 7.3 6.5 6.2 4.5 4 .3 4.0 1.5 2.2 T a b le 11. W i n t er s u r v i v al as i n f l u e n c ed by s n o w m o ld a nd w i n t e r- k i ll u n d er f o r a ge plot c o n d i t i o n s, U. S. P a s t u re L a b- o r a t o r y, U n i v e r s i ty P a r k, P a. 1971-72. V a r i e ty P e n n f i ne " N o r l ea " T e t r a p l o id " Linn " P e n n f i ne " " N o r l ea " 11 » T e t r a p l o id " " " Linn " " " N i t r o g en K g / Ha % S u r v i v al A p r il 19 71 50 100 50 100 50 100 50 100 0 50 100 200 0 50 100 200 0 50 1 00 200 0 50 100 200 100 100 9 0 90 70 100 35 70 A p r il 19 72 99 67 43 50 94 74 55 46 43 13 11 1 54 9 4 3 T a b le 12. R y e g r a ss t u rf e v a l u a t i on -- f l u o r e s c e n c e, v e r t i c al g r o w t h, and leaf width, U n i v e r s i ty P a r k, P a. V a r i e ty % f l u o r e s c e n ce S e e d l i ng v e r t i c al g r o w th Leaf width, mm Leaf 1 Leaf 2 0 0 5.0 0 0 3.5 0 4.0 9 1 .0 P e n n f i ne M a n h a t t an N K- 100 P e lo N o r l ea B r a n b a n t ia C o m bi L i nn A n n u al LSD .05 40 50 80 80 60 70 80 80 100 15 2.5 2.7 2.9 3. 1 3. 1 3.6 3.3 3.8 5.6 0.5 2.6 2.7 2.9 3.1 3.1 3.6 3.4 3.8 5.8 0.5 T a b le 13. R y e g r a ss o v e r s e e d i ng e v a l u a t i o n, S t a te C o l l e g e, M i s s- i s s i p p i, 1969-70. W i n t er t u rf q u a l i ty e v a l u a t i o n s* V a r i e ty 1 1 / 12 1 2 / 18 l /l 3 2 / 11 3 / 26 4 /9 5 /5 6 /5 P e n n f i ne M a n h a t t an M e d a l i st II A n n u al M a g n o l ia 6.5 7.2 6.6 7. 7 8.2 6.7 7.0 6.0 7.5 6.5 6.7 7.2 6.7 7.0 6.5 5.2 6.2 4.0 2.7 3.0 8.7 7.7 5.5 5.2 5.5 6.7 6.7 4 .5 6.0 4.7 4.0 1.7 1.7 1.5 1.0 7.0 6.5 4 .5 3.5 3.0 * R a t i ng s c a l e: 1 to 10; 10 i b e s t, 5 : a c c e p t a b le t u r f. T a b le 14. R y e g r a ss o v e r s e e d i ng e v a l u a t i o n, C o l l e ge S t a t i o n, T e x a s, 1969-70. D a ys to Avg. p u t t i ng e m e r g e n ce C o l or s u r f a c e* A v e r a ge t r a n s i t i o n* P oa a n n ua c o m p e t i t i on P e n n f i ne M a n h a t t an N K- 100 L i nn P e lo M e d a l i st II A n n u al 5 -6 7 5 5 5 5 3 m ed to d a rk m ed light to m ed light light to m ed light to m ed light 1.2 1.1 1.6 1.9 1.9 1.3 2 .4 2.0 2.0 3.1 3.1 2 .4 3.0 3.3 good f a ir good good good good good * S c a l e: 1 to 5; 1 - e x c e l l e n t, 5 = u n a c c e p t a b l e. S E E K I NG D W A R F N E SS IN B L U E G R A SS W i l l i am H. D a n i el D e p a r t m e nt of Agronomy- P u r d ue University- L a f a y e t t e, I n d i a na 47907 I n t r o d u c t i on In looking at t he b l u e g r a ss b r e e d i n g, s o me of t he i n t e r e s t i ng q u e s t i o ns a r e: How i n f r e q u e n t ly could one m o w? How d w a rf would you like t u rf to b e? And, of c o u r s e, t he m o st b a s ic q u e s- t i on -- h ow do you get i t? A n o t h er q u e s t i on i s, of c o u r s e, s p r e a d. We c an get s p r e ad of 4 to 8 t i m es M e r i o n, t he s t a n d a rd in t e s t s. h a ve a n e ed f or v i g o r; t h e r e f o r e, we n e ed t h i n gs t h at p r o d u ce a lot of s p r e a d, f i l l - in and r e c o v e ry f or a t h l e t i c s, p a r k s, s i d e s, p o or n u t r i t i on a r e as and s i m i l ar p l a c e s. One w ay to m e a s u re s p r e ad is by i n d i v i d u al c l o ne s p r e ad as i n c h e s, m e t e rs a nd r a t i n g s. In c e r t a in p l a c es we r o a d- S e e d l i ng V i g or A n o t h er t h i ng we h a ve t a l k ed about a g r e at d e al is s e e d l i ng is s l o w er t h an m a n y. P a rk c l a i m ed v i g o r. M e r i on b l u e g r a ss s e e d l i ng v i g o r. We a l so k n ow b l u e g r a s s es v a ry in how t h e ir a n g le out a l o ng t he s t e m s. We h a ve t e n d ed to t u rn o ur b a c ks on t he c o m m on t y p es and t h o se t h at t e nd to h a ve a n a r r ow leaf a n g l e, as w e ll as i n i t i al leaf e m e r g e n ce h i gh a b o ve t he g r o u n d. l e a v es In 19 62 we b e g an to n o t i ce d w a r f n e ss in c e r t a in p l a n t s. We h a ve c o l l e c t ed f or t en y e a r s. T h e re a re o v er 100 v a r i a t i o ns w h i ch c a me f r om one p l a nt c h o s en in 19 62. u n m o w ed f or one y e ar v a r i ed f r om 4 to 20 i n c h e s. We w e re p l e a s ed with t he s p r e ad of s o me of t he p l a n t s. S e v e r al d w a r fs p r o d u ce l e ss s e e d h e a ds down v e ry low, q u i te l a te in s e ed p r o d u c t i o n. In t he g r e e n h o u s e, l e a v es Leaf H e i g ht We h a ve k e pt a b o ut 50 of t he b e t t er d w a r fs w i th leaf h e i g ht of l e ss t h an 3 i n c h e s. T h ey a re u s u a l ly s l ow in s p r e ad and m a ny of t h em t e n d ed to f a i l. You w a nt s o m e t h i ng t h at h as r h i z o me t h at d o es e m e r g e, f i ll out, k e e ps c o m i n g, and h as t he a b i l i ty to r e j u v e n a te i t s e l f. S e e d h e a ds h a ve b e en v e ry s p a r se on m o st d w a r f s, p a r t i c u l a r ly d u r i ng t he f i r st y e ar of p r o d u c t i o n, with low s e e d h e ad h e i g ht - 20 c m ., as an e x a m p l e. D u r i ng t he s e c o nd y e a r, n e ar n o r m al s e ed w as h a r v e s t e d. A m o ng a ll b l u e g r a s s es in t e s t, s e e d h e ad h e i g h ts v a ry f r om 30 i n c h es down to 5 i n c h e s. U s u a l ly s e e d h e ad h e i g ht is about t w i ce leaf s p r e a d, l e n g t h. M a ny of t he g r a ss v a r i e t i es n ow on t he m a r k e t, or to c o me on t he m a r k e t, a re n o t ed f or t he a b i l i ty to p r o d u ce l e a v es c l o se to t he g r o u n d. T h is is one of t he t a r g e ts of d w a rf v a r i e t i e s. T he p l a nt w i th t he t e n d e n cy to d e v e l op l e a v es c l o se to t he r e s u l t ed in m o re t i l l e rs -- m o re l e a v es t y p e s. c o m p a r ed to m o re c o m m on g r o u nd h as S u m m a ry B r i e f l y, we h a ve m a de s e v e r al s e l e c t i o n s, and we a re i n t e r- e s t ed in d w a r f n e s s. O ur q u e s t i on i s: How d w a rf c an it b e? And, on t op of it - how do you s e r ve t he s e ed g r o w er and t he s e ed p r o- c e s s or A - 20 or A - 3 4, t h at is s i m p l er t h an t r y i ng to t a ke it a ll t h r o u gh s e ed f i e ld of o ur n o r m al p r o c e d u r e. D i s e a se r e s i s t a n ce h as a l- w a ys b e en i m p o r t a nt w h a t e v er t he v a r i e t y. if you go t h is r o u t e? If one g o es v e g e t a t i v e, as with W a r r e n 's t he D o es t he d w a r f n e ss fit into t he f u t u r e? If so, w i ll it be v e g e t a t i v e? How d w a rf c an we go as we w o rk t o w a r ds b l e n d i ng w i ll be an i n t e r e s t i ng q u e s t i o n. C e r t a i n ly it w i ll t a ke a lot of f i n e s se in m a n a g e m e n t. T HE O R I E N T A T I O NS OF T HE R H I Z O M ES OF K E N T U C KY B L U E G R A SS J o hn E. F i s h er R e s e a r ch S t a t i on C a n a da D e p a r t m e nt of A g r i c u l t u re O t t a w a, C a n a da KIA O C6 I n t r o d u c t i on T he p o p u l ar c o n c e pt of t he r h i z o me (a l ax u n d e r g r o u nd s t em g r o w i ng t h r o u gh t he s o il u n t il i ts t ip e v e n t u a l ly t u r ns up to b e c o me a n ew p l a n t) i m p l i es a p a r t i c u l a r ly d i s o r i e n t ed and d i s o r g a n i z ed s t r u c t u re w a n d e r i ng a i m l e s s ly t h r o u gh t he s o il without r e al o r i e n t a t i o ns or c o n t r o l s, a p a rt f r om t h o se by i m m o v a b le o b j e c ts in t he s o i l. T h is m i s c o n c e p t i on is not m e r e ly due to l a ck of k n o w l e d ge by t he l a y m a n, but is by t he d e s c r i p t i on p r e s e n t ed in m o st b o t a n i c al t e x t b o o ks and in one of t he m o st p r o m i n e nt m a n u a ls of t he g r a s s e s. H o w- e v e r, s t u d i es h a ve s h o wn t h at t he r h i z o m es of g r a s s es a re u n d er p r o n o u n c ed o r i e n t a t i o n al c o n t r ol f r om t he t i me of t h e ir i n i t i a t i on as r h i z o me b u d s, u n t il t h e ir t i ps u l t i m a t e ly e m e r ge f r om t he s o il and b e c o me a e r i al ( l e a f y) s h o o t s. i m p o s ed i m p l i ed F or t he f i r st t i m e, t he o r i e n t a t i o ns of t he K e n t u c ky b l u e- g r a ss r h i z o me d u r i ng i ts l i fe h i s t o ry w i ll be d e s c r i b ed in t h e ir c h r o n o l o g i c al and b i o l o g i c al s e q u e n c e. S o me of t h e se o b s e r v a- t i o ns h a ve b e en p r e s e n t ed p r e v i o u s ly ( F i s h e r, 1964, 1965, 1966a, 19 66b, 19 72), w h i le o t h e rs h a ve not. T he O r i g in and I n i t i al O r i e n t a t i on of R h i z o me B u ds G r a s s es a re not t r u ly b i l a t e r a l ly s y m m e t r i c a l. T he point of a t t a c h m e nt of e a ch leaf d e v i a t es s l i g h t ly f r om t he p l a ne of b i l a t e r al s y m m e t r y. One leaf m ay be d i s p l a c ed in one d i r e c- t i o n, t he next in t he o p p o s i t e, or s e v e r al c o n s e c u t i ve l e a v es m ay be d i s p l a c ed in t he s a me d i r e c t i o n. T h is is b e l i e v ed to r e f l e ct t h e ir p h y t o g e n e t ic r e l a t i o n s h ip to a p r i m i t i ve t h r ee p r o g e n i t o r. We h a ve f o u nd no m e a ns of p r e d i c t i ng t he d e v i a- t i on of f u t u re l e a v es not yet i n i t i a t e d. H o w e v e r, o n ce t he f i r st c e ll d i v i s i on in t he i n i t i a t i on of a n ew leaf h as t a k en p l a c e, t he d e v i a t i on of t h at leaf f r om t he p l a ne of s y m m e t ry h as b e en d e t e r m i n ed and t h is d e v i a t i on c an be o b s e r v ed m i c r o s c o p i c a l l y. r a n k ed t i e r s, ( F i g. 1; F in F i g. 2). is e l l i p t i c al in c r o ss s e c t i o n, not c i r c u l a r. In K e n t u c ky b l u e g r a ss t h is h as p r o v ed to be s i m p l er t h a n, f or e x a m p l e, t he c e r e al g r a s s e s. T he a p i c al m e r i s t em of b l u e- g r a ss S u r m o u n t i ng t he a p ex and e x t e n d i ng down both f l a n ks is a p a ir of c e ll o ne on e a ch s i de of t he p l a ne of s y m m e t ry W h en a n ew leaf is about to be i n i t i a t e d, a c e ll u n d e r g o es a p e r i c l i n al c e ll d i v i s i o n, i n c r e a s i ng t he g i r th of t he a p ex c r o ss s e c t i on at t h at point ( a n g le B, F ig 2). D i v i s i o ns s p r e ad a r o u nd t he a p ex in both d i r e c t i o n s, m e e t i ng at a point d i s p l a c ed f r om t he l i ne of s y m m e t ry on t he o p p o s i te s i de of t he a p ex ( a n g le A, F i g. 2). T h u s, one s i de of t he leaf p r i m o r d i um is of g r e a t er p a r t i al c i r c u m f e r e n ce t h an t he o t h e r. T h is d i s p a r i ty in s i ze v i s i b le in t he m a t u re leaf w h e re " h a l f" of t he b l a de is s l i g h t ly w i d er t h an t he o t h e r. It is t h is d e v i a t i on f r om t he line of s y m- m e t ry of t he i n i t i a t i ng leaf t h at d e t e r m i n es t he d e v i a t i on f r om s y m m e t ry of t he p o s i t i o n i ng of t he a x i l l a ry bud f r om w h i ch a r h i z o me ( or a t i l l e r) d e v e l o p s. is T he bud p r i m o r d i um a r i s es f r om c e ll d i v i s i o ns d e ep w i t h in It is p o- t he b a se of t he leaf p r i m o r d i um ( F i g. 3). T h e r e f o r e, t he bud d e v e l o ps f r om t i s s ue giving r i se to t he leaf a b o v e. s i t i o n e d, c i r c u m f e r e n t i a l l y, at t he point w h e re t he t wo s e m i- c i r c l es of t he d e v e l o p i ng leaf p r i m o r d i um c a me t o g e t h e r; i s, on t he s i de o p p o s i te t he f u t u re p o s i t i on of t he m i d - r ib of t he leaf a b o ve ( F i g. 2). T he c o n c e pt t h at a bud a r i s es in t he a x il of t he leaf i m m e d i a t e ly b e l ow t he bud is c o n v e n i e nt g e o- g r a p h i c a l l y, but b i o l o g i c a l ly t he bud is m o st c l o s e ly r e l a t ed to t he leaf a b o v e, w h o se b l a de w i ll be on t he o p p o s i te s i de to t he bud ( S h a r m a n, 1942, 1945, 1947; E t t e r, 1951). T h is unit of s t r u c t u r e, w h i ch u l t i m a t e ly c o n s i s ts of t he leaf a b o v e, a s e c- t i on of s t em ( t he i n t e r n o d e ), and a bud at t he b a se of t he i n t e r- n o de on t he s i de o p p o s i te t he m i d - r ib of t he leaf a b o v e, h as t h at F i g u re 1. T op v i ew of t he living a p ex of K e n t u c ky b l u e- g r a s s. T he l i n es s h ow t he p l a ne of b i l a t t e r al s y m m e t ry and point to t he p a i r ed t i e rs of c e l l s, one t i er on e i t h er s i de of t he p l a ne of s y m m e t r y. F i g u re 2. D i a g r a m m a t ic r e p r e s e n t a t i on of a b l u e g r a ss s e r i al s h o o t, (A) a n g u l ar d e v i a t i on of t he leaf v i e w ed f r om a b o v e: s h e a th f r om b i l a t e r al s y m m e t r y; (B) a n g u l ar d e v i a t i on of m e d i an s t r a nd f r om b i l a t e r al s y m m e t r y; (C) a p i c al m e r i s t e m; (D) leaf s h e a t h; ( E) a r r o ws r e p r e s e nt r a t es of d e v e l o p m e nt of a f o l i ar p r i m o r d i um a r o u nd t he a p i c al m e r i s t e m; ( F) p a i r ed t i e rs of c e l ls s t r a d d l i ng t he p l a ne of s y m m e t ry of t he a p i c al m e r i s t e m; m e r i s t e m; (I) y o u n g e st c a t a p h y l l, and (J) f i r st t r ue c a t a p h y ll of bud. F r om F i s h e r, J. E ., Can. J. B o t a ny 50: 1972. (G) bud p r o p h y l l; (H) bud a p i c al 7 4 3 - 7 5 0, F i g u re 3. L o n g i t u d i n al s e c t i on on t he p l a ne of s y m m e t ry of a (A) a p i c al m e r i s t e m; (B) bud p r i m o r- r h i z o me t i p: d i um d e v e l o p i ng f r om t he b a se of a leaf p r i m o r d i u m; ( Li , L2> L 3) leaf p r i m o r d ia in o r d er of a g e. E a ch p r i m o r d i um e x c e pt t he y o u n g e s t, Li , c o n s i s ts of two p o r t i o ns w h en v i e w ed in l o n g i t u d i n al s e c t i o n. b e en c a l l ed a p h y t o m er ( E v a n s, 1949; E t t e r, 19 51; F i s h e r, 1965, 19 72). T h u s, t he p o s i t i on of t he a x i l l a ry bud w as d e t e r m i n ed by t he p o s i t i on of t he f i r st c e ll d i v i s i on in g a ve r i se to t he leaf p r i m o r d i u m. t he a p i c al m e r i s t em t h at In t he d e v e l o p m e nt of t he bud, t he b i o l o g i c a l ly c o n t r o l l ed o r i e n t a t i on of o r g a ns i n i t i a t ed o r i g i n a l ly by t he a c t i v i ty of a s i n g le m e r i s t em c e ll is c o n t i n u e d. T he bud p r o d u c es f o l i ar o r- g a n s, t he f i r st of w h i ch is t he t w o - v e i n ed p r o p h y l l. w i th i ts l a r g er s i de n e xt to t he m a in shoot f r om w h i ch t he bud a r o se ( F i g. 2). T he bud c a t a p h y l ls ( r e d u c ed l e a v es with a t r ue m i d - r i b, c o m m o n ly c a l l ed bud s c a l e s) a re o r i e n t ed with t h e ir p l a ne of s y m m e t ry a p p r o x i m a t e ly at r i g ht a n g l es ' to t h at of t he m a in s h o ot (J in F i g. 2). It is o r i e n t ed F i g u r es 4, 5. C r o ss s e c t i o ns of r h i z o me b u ds s h o w i ng r o t a t i on of t he p l a ne of s y m m e t ry of t he bud a p e x. T he b l a ck l i n es a re p o s i t i o n ed on t he p l a n es of s y m m e t ry of t he a p ex of e a ch bud. F r om F i s h e r, J. E ., Can. J. B o t a ny 50: 7 4 3 - 7 5 0, 1972. N o n - g e o t r o p ic O r i e n t a t i o ns D u r i ng t he I n i t i al G r o w th P h a se of R h i z o me B u ds A f t er a r h i z o me bud h as f o r m e d, it m ay r e m a in in a d o r m a nt In f a c t, r h i z o me buds on e x- c o n d i t i on f or long p e r i o ds of t i m e. i s t i ng r h i z o m es o f t en n e v er d e v e l op b e y o nd t he f i ve or s ix c a t a- p h y ll s t a ge ( E t t e r, 1951, 1953). If a c t i ve g r o w th of a bud c o m m e n c e s, t he p r o p h y ll and t he o l d e st one or t wo c a t a p h y l ls a re not u s u a l ly i n v o l v e d. T h ey a re n o r m a l ly r u p t u r ed by t he e x p a n d i ng c a t a p h y l ls w i t h in and r e m a in as d ry t a t t e r ed s c a l es at t he b a se of t he bud ( F i g. 4, 5, 6). T he r h i z o me bud d e v e l o p i ng f r om t he b a se of an a e r i al plant t u r ns d o w n w a r ds p r e p a r a t o ry to a c t i ve e l o n g a t i ve g r o w th into t he s o i l. T h is r e a c t i on is a p p a r e n t ly a p o l a r i ty r e a c t i o n, r e l a t ed t he f o re and aft a x is of t he shoot f r om w h i ch t he bud a r o s e, and not c a u s ed by s u s c e p t i b i l i ty of t he bud to t he f o r ce of g r a v i ty ( F i g. 6, 7). R h i z o me buds on p l a n ts g r o w i ng on t he r o t a t i ng c l i n o s t at r o t a t i ng at 10 r e v o l u t i o ns p er h o ur ( g r a v i ty s t i m u l a t i on b e i ng m u l t i d i r e c t i o n al r a t h er t h an u n i d i r e c t i o n a l) o r i e n t ed t h e m- s e l v es w i th r e s p e ct to t he f o re and aft a x is of t he s t em f r om w h i ch t h ey a r o s e. T h ey did not a p p e ar to be a f f e c t ed by g r a v i t y. L a t e r, g r a v i t a t i o n a l ly s e n s i t i ve o r i e n t a t i o ns w e re s e v e r ly d i s o r g a n i z ed by g r o w th on t he c l i n o s t a t. to T he i n i t i al b e n d i ng a p p e a rs to be a f u n c t i on of t he c a t a p h y l l s. C a t a p h y ll v e i ns t h i c k en and d e v e l op s c h l e r e n c h y m a t o us c e ll w a l ls m o re r a p i d ly on t he s i de a w ay f r om t he s t em ( o u t er or a b a x i al s i d e ), h e n ce a c t i ve g r o w th of t he s i de of t he c a t a p h y l ls next to t he s t em ( a d a x i a l ), c a u s es b e n d i ng of r h i z o m e s. As t he c a t a p h y l ls in s u c c e s s i on f o rm a t e l e s c o p i ng t u be t h r o u gh w h i ch t he young r h i- z o me bud s t em g r o w s, t he r h i z o me s t em is t h e r e f o re d i r e c t ed a d o w n w a r ds d i r e c t i on by d i f f e r e n t i al c a t a p h y ll g r o w t h, and not by a c t i ve b e n d i ng of i ts own t r ue s t e m. E x c i s ed s t e ms of young r h i z o me b u ds a re r e l a t i v e ly s t r u c t u r e l e ss and v e ry f l a c i d. into T he p l a ne of s y m m e t ry of t he a p ex of t he young bud w as o r i- g i n a l l y, a p p r o x i m a t e ly at r i g ht a n g l es to t h at of t he m a in shoot ( F i g. 2). If t h is c o n t i n u e d, t he p l a ne would u l t i m a t e ly be p a r a l l el to t he s o il s u r f a c e, or m o re c o r r e c t l y, at r i g ht a n g l es to a p l u mb l i n e. H o w e v e r, t he p l a ne of s y m m e t ry of t he r h i z o me a p ex d u r i ng a c t i ve e l o n g a t i on is at r i g ht a n g l es to t he s o il s u r f a c e, h e n ce p a r- a l l el to a p l u mb l i n e. T h is r o t a t i on of s y m m e t ry of t he r h i z o me a p ex t a k es p l a ce d u r i ng t he f o r m a t i on of t he f i r st 7 to 10 c a t a- p h y l l s, and p r i or to t he m o st a c t i ve e l o n g a t i on p h a se ( F i g. 4, 5, 8). T h is s h i ft is at l e a st p a r t i a l ly i n d e p e n d e nt of g r a v i t y, s i n ce buds F i g u re 6. T he i n i t i al d o w n w a r ds b e n d i ng of a young r h i z o me bud. Note t he p r o m i n e nt v e i n s. F i g u re 7. F r o nt v i ew of t he bud in F i g u re 6, s h o w i ng t he b e g i n n i ng of l a t e r al b e n d i ng t o w a r ds t he r i g h t. on t he p l a n ts on t he c l i n o s t at s h ow r o t a t i on of t he p l a ne of s y m m e t r y. H o w e v e r, t h ey do not c e a se t he r o t a t i on of t he p l a ne of s y m m e t ry at a p a r t i c u l ar a n g l e, e i t h er to t he s o il s u r f a ce or to t he m a in p l a nt a x is and no two s e em to c e a se at t he s a me a n g l e. T he g r a v i t a t i on s u s c e p t i b i l i ty s e e ms to o c c ur t o w a r ds t he l a t t er s t a g es of r o t a t i on of t he p l a ne of s y m m e t ry w h en i n t e r n o de e l o n g a t i on of t he young r h i z o me s t em b e c o m es a c c e l e r a t e d. F i g u re 8. D i a g r a m m a t ic i l l u s t r a t i on of t he r o t a t i on of s y m- m e t ry in t he d e v e l o p i ng r h i z o me of K e n t u c ky b l u e- (A) t op view, t he a r r o ws r e p r e s e nt t he p l a ne g r a s s: of s y m m e t ry at e a ch l o c a t i o n. T he m a in shoot is at t he r i g ht s i d e: r h i z o m e. E a ch c a t a p h y ll is n u m b e r e d. T he c i r c l es b e n e a th r e p r e s e nt c r o s s - s e c t i o ns at e a ch of t he n u m b e r ed c a t a p h y l l s. T he a r r o ws a b o ve e a ch c i r c le s h ow t he p l a ne of s y m m e t ry of t he m a in s t em f r om w h i ch t he r h i z o me a r o s e. (B) s i de v i ew of t he d e v e l o p i ng G e o t r o p ic O r i e n t a t i o ns D u r i ng t he A c t i ve E l o n g a t i ve P h a s es of R h i z o me G r o w th T he young d e v e l o p i ng r h i z o me a p p e a rs to b e c o me s e n s i t i ve r h i z o me to g r a v i t a t i o n al f o r c es w h en a c t i ve i n t e r n o de e l o n g a t i on b e c o m es p r o n o u n c e d. T he g r o w th p h a se d u r i ng w h i ch t he young is e x t e n d i ng d e e p ly into t he s o il at an o b l i q ue a n g le to a p l u mb l i ne (a p l a g i o g e o t r o p ic r e s p o n s e ), I h a ve t e r m ed t he p r i m a ry s t a ge ( F i s h e r, 19 65). A f t er t he c o n t a i n e rs in w h i ch t he p l a n ts w e re g r o wn w e re t i l t ed by a p r e d e t e r m i n ed n u m b er of d e g r e e s, t he r h i z o m es r e o r i e n t ed t h e m s e l v es so as to a s s u me t h e ir f o r- m er a n g le to t he p l u mb l i n e. T he c o n t o ur of t he s o il s u r f a ce d o es not a p p e ar to i n f l u e n ce t he o r i e n t a t i on of b l u e g r a ss D u r i ng t h is p l a g i o g e o t r o p ic s t a ge t he c a t a p h y l ls a re q u i te d i s t i n c t i v e. V i e w ed in t h e ir e n t i r e t y, t h ey a re d i s t i n c t ly c o n e - s h a p e d. s e c t i o n, t h ey a re e l l i p t i c al to a l m o st c i r c u l ar and m o re h e a v i ly w a l l ed on t he s i de of t he m a in v e i n. T h ey do not h a ve a t u d i n al i n d e nt or i n v a g i n a t i on on t he t h i n n er s i d e. T he t ip of t he young c a t a p h y ll o f t en h as a s m a ll t r i a n g u l ar p o r e - s h a p ed o p e n i ng c l o se to, but s l i g h t ly b e n e a th i ts tip. O c c a s i o n a l ly t h e re is no o p e n i ng e v i d e n t. r h i z o m e s. In c r o ss l o n g i- T he s e c o n d a ry s t a ge b e g i ns w i th a c h a n ge in t he s h a pe and f o rm of t he n ew c a t a p h y l l s. T h ey b e c o me m a r k e d ly e l l i p t i c al and h a ve an i n d e n t a t i on or i n v a g i n a t i on r u n n i ng t he f u ll l e n g th of t he c a t a p h y ll on t he t h i n - w a l l ed s i de o p p o s i te to t he m a in vein. In o v e r a ll view, t h ey a re d i s t i n c t ly s w o r d - s h a p e d. T he p o re at t he t ip b e c o m es a d e f i n i te s l i t. T h en a m a j or c h a n ge in t he g e o- t r o p ic r e s p o n se t a k es p l a ce as t he tip, w h i ch h ad b e en g r o w i ng d o w n w a r d s, r a p i d ly s h i f ts i ts d i r e c t i on into t he h o r i z o n t al p l a n e. U n d er i d e al m i d s u m m er c o n d i t i o n s, t he s e c o n d a ry s t a ge is l o n g e st of any. t he D u r i ng t he f i n al or t e r t i a ry s t a ge a s m a l l, i n c o m p l e t e, m e m- b r a n e o us l i g u le f o r ms at t he b a se of t he a p i c al slit in t he f i r st c a t a p h y l l. T he f i r st s i g ns of t he l a r g e, t h i n - w a l l ed c e l ls b e l i e v ed to be i n v o l v ed in leaf c l o s u re in m a t u re a b o v e - g r o u nd l e a v es c an j u st be o b s e r v ed a b o ve t he l i g u le on e i t h er s i de of t he m a in vein. E a ch of t he next f i ve s u c e s s i ve c a t a p h y l ls b e g i ns to d e v e l op in m a t u re l e a v e s, c a u s es t he b l a de t he c o l l ar l e a f - l i ke c h a r a c t e r i s t i c s. H o w e v e r, c e r t a in t h a t, a w ay f r om t he a x is of t he s h e a th d o es not d e v e l op in s t a ge c a t a p h y l l s. D u r i ng t he f i n al s t a ge in g e o t r o p ic t he r h i z o me t ip b e g i ns to t u rn u p w a r d, d e s c r i b i ng a long a rc t h r o u gh t he s o il u n t il it f i n a l ly e m e r g es and c h a n g es to a n ew a e r i al s h o o t. T h is g e o t r o p ic r e s p o n se h as b e en c a l l ed n e g a t i ve o r t h o- g e o t r o p i s m, now f r e q u e n t ly a b b r e v i a t ed to a p o g e o t r o p i s m. to b e nd o b l i q u e ly t e r t i a ry r e g i on r e s p o n s e, W h en t he t ip of t he r h i z o me p e n e t r a t es t he s o il s u r f a ce t he f i r st t wo or t h r ee o r g a ns a re not t r ue l e a v e s, but m e r e ly g r e en c a t a p h y l ls of t he t e r t i a ry s t a g e. T h ey o r i g i n a t ed f r om t he a p ex b e f o re t he r h i z o me h ad e m e r g ed f r om t he s o i l. T h e r e a f t e r, n o r- m al a e r i al l e a v es a re p r o d u c e d. In e a ch of t he t h r ee s t a g es - - p r i m a r y, s e c o n d a r y, m o r p h o l o g i c al f o rm and s t r u c t u re of t he c a t a p h y l ls is l i n k ed with a s p e c i f ic p h y s i o l o g i c al r e s p o n se to g r a v i ty to s u ch an e x t e nt t h at ojie c an be u s ed as an a c c u r a te i n d i c a t or of t he o t h er ( F i s h e r, 1965, 1972). t e r t i a r y -- i n t i m a t e ly C i r c u m n u t a t i o n al G r o w th M o v e m e n ts of R h i z o m es R h i z o m es c an p e n e t r a te o b j e c ts in t he s o i l, s u ch as wood, a s p h a l t, h e a vy clay, r o o ts and t u b e rs of o t h er s p e c i e s, and a l m o st a n y t h i ng e l se in t h e ir p a t h s, e x c e pt s t o n es or m e t a l s. T h ey do so by m e a ns of c ir c u m n u t a t i o n al g r o w th of t he i n d i v i d u al c a t a p h y l l s. C i r c u m n u t a t i on h as b e en d e s c r i b ed ( L a r s o n, 19 62) as t he s p i r al p a t h w ay in s p a ce d e s c r i b ed by t he t ip of a s t em d u r i ng g r o w t h. A p p a r e n t ly m o st s t e ms exhibit s o me d e g r ee of c i r c u m n u t a t i o n. T r ue c i r c u m n u t a t i on is not a t o r s i o n al t w i s t i n g, s u ch as e x h i b i t ed by m o r n i ng g l o ry v i n e s; h e n c e, is not a n a l a g o us with a s c r e w. To d e m o n s t r a te t he m o v e m e n ts of a r h i z o me tip, an a p p a r a t us w as c o n s t r u c t ed w h i ch c o n s i s t ed of a s h o r t, p i v o t e d, b a l a n ce a r m, h a v i ng a c o ne at one end into w h i ch a r h i z o me t ip w as guided. At t he o t h er end w as an i n c l i n ed m i r r or onto w h i ch a c o l l i m a t ed light b e am w as d i r e c t e d, t he r e f l e c t i on b e i ng p r o j e c t ed onto t he d i s t a nt w a ll of a d a r k e n ed r o om ( F i s h e r, 19 64). A c i r c u l a r, c l o c k w i s e, 2 4 - h o ur r o t a t i on of t he r h i z o me t ip w as o b s e r v ed at 1,000 X m a g n i f i c a t i o n. T he a c t u al d i a m e t er of r e v o l u t i on w as b e t w e en 0.5 and 3.0 m m. A m i n o r, c i r c u l a r, c l o c k w i se r e v o l u t i on of one to two h o ur p e r i o d s, s u p e r i m p o s ed on t he m a j or r e v o l u t i o n, w as d e t e r m i n ed at 10,000 X m a g n i f i c a t i o n. T he a c t u al d i a m e t er of r e v o l u t i on w as b e t w e en 0.001 and 0.010 m m. In soil, t he m a g n i t u de of t h e se r o t a t i o ns would p r o b a b ly be m u ch l e ss due to t he p h y s i c al d a m p i ng a c t i on of t he soil. T h e se r o t a- t i o n al m o v e m e n ts u n d o u b t e d ly aid t he r h i z o me in p e n e t r a t i on of t he soil. H o r i z o n t al O r i e n t a t i on of R h i z o me B u ds i ts p l a ne of b i l a t e r al s y m m e t ry is p a r a l l el to t he If a plug of b l u e g r a ss sod is p l a n t ed in a v e g e t a t i o n - f r ee a r ea of soil, n ew r h i z o m e s, w h en p r o d u c e d, w i ll g r ow o u t w a rd m u ch like t he s p o k es of a w h e e l. U l t i m a t e ly t h ey t u rn up to p r o d u ce n ew a e r i al or l e a fy s h o o t s. W h en t he d a u g h t er plant h as e m e r g e d, d i r e c t i on in w h i ch t he r h i z o me h as b e en g r o w i n g. T h e r e f o r e, r h i z o me b u ds p r o d u c ed by t he d a u g h t er plant w i ll be p o s i t i o n ed e i t h er d i r e c t ly a w ay f r om t he o r i g i n al p a r e nt p l a n t, or d i r e c t ly t o w a r ds it ( F i g. 9). S e r e b r y a k o va ( 19 68) h as s t a t ed t h at t he bud p o s i t i o n ed t o w a r ds t he p a r e nt p r o b a b ly n e v er d e v e l o ps s i n ce one n e v er s e es a r h i z o me g r o w i ng b a ck t o w a r ds t he p a r e nt p l a nt w h en t he p a r e nt is i s o l a t ed and not in a d e n se sod. I h a ve f o u nd t h at t h e se b u ds do d e v e l op but t he r h i z o m es d e- v e l o p i ng f r om t h em b e nd t i g h t ly a r o u nd t he s t em of t he d a u g h t er p l a nt u n t il t h e y, l i ke t he b u ds on t he o t h er s i de of t he d a u g h t er p l a n t, a re g r o w i ng a w ay f r om t he o r i g i n al p a r e nt ( F i g. 10, 11). T h is h i g h ly o r i e n t ed b e n d i ng is a c c o m p l i s h ed t h r o u gh d i f f e r e n t i al g r o w th r e s p o n s es in t he c a t a p h y l ls of t he r h i z o m e. One s i de of t he bud g r o ws at a m u ch h i g h er r a te t h an t he o t h er ( F i g. 6, 7). T h is b e n d i ng a r o u nd t he a e r i al shoot r e s e m b l es c l o s e ly t he b e n d i ng d o w n w a r ds of t he young r h i z o me bud, as o u t l i n ed in t he p r e c e d i ng p o r t i on of t h is r e p o r t. t ip e m e r g es f r om t he soil, t h is o r i e n t a t i on of s u b s e q u e nt z o m es d o es not o c c u r. s o il p r i or to s e v e r i ng and h as f o r m ed a n ew a e r i al shoot, t he o r i e n- t a t i on r e m a i ns f or m o re t h an f o ur w e e k s. T he o r i e n t a t i on a w ay If t he r h i z o me t ip h as e m e r g ed f r om r h i- If t he r h i z o me is s e v e r ed b e f o re t he r h i z o me t he F i g u re 9. D i a g r a m m a t ic r e p r e s e n t a t i on of t he m o v e m e nt of r h i z o m es a w ay f r om t he o l d er p l a n ts in a r h i z o me - a e r i al shoot s e r i e s. R h i z o me b u ds t h at a r i se f a c i ng t he r h i z o me - a e r i al shoot s e r i es bend a r o u nd t he b a se of t he shoot and m o ve o u t w a r ds plant in s e r i e s; ( P2 ) y o u n g er p l a nt in s e r i e s; z o me c o n n e c t i ng Pi and P2 ; ( R 2) young r h i z o m es a r i s i ng f r om t he b a se of P2 . into n o n - i n h a b i t ed soil. (R) r h i- (F\ ) O l d er f r om t he o r i g i n al p l a nt d o es not a p p e ar to be t he r e s u lt of t o x ic or g r o w t h - r e g u l a t i ng s u b s t a n ce s e c r e a t ed into t he s o il by t he p a r e n t. N e w ly e m e r g ed d a u g h t er p l a n t s, if l o c a t i on but r o t a t ed by 180°, w i ll p r o d u ce r h i z o m es t h at will g r ow d i r e c t ly t h r o u gh t he p a r e nt p l a nt without a ny a p p a r e nt i n h i b i t i on of g r o w t h. A p p a r e n t ly t he i n n er buds b e nd due to s o me p o l ar t y pe s t i m u l us f r om t he d a u g h t er p l a nt a n d / or f r om t he r h i z o me f r om w h i ch t he d a u g h t er a r o s e. t r a n s p l a n t ed in t h e ir s a me T h u s, t he r h i z o m es d e v e l o p ed by e a ch s u c c e e d i ng plant o r i e nt t h e m s e l v es with r e s p e ct to t he p r e c e d i ng p l a nt in e a ch t r a in ( F i g. 9), m a k i ng t he b l u e g r a ss p l a nt a p o t e nt i n v a d er of n ew a r e as of soil. B e s i d es t he r h i z o m es i n i t i a t ed by t he p a r e nt and by t he d a u g h t er p l a n t s, c e r t a in d o r m a nt b u ds along t he r h i z o m es r h i z o me F i g u re 10. P r o m i n e nt l a t e r al b e n d i ng of a r h i z o me bud. F i g u re 11. T he b e n d i ng a w ay f r om t he p a r e nt r h i z o me (E), and a r o u nd t he a e r i al shoot ( F) of a r h i z o me bud (A) and a d e v e l o p ed r h i z o me (B). R h i z o me (C) and r h i z o me bud (D) h a ve d e v e l o p ed f r om t he o p p o s i te s i de of t he a e r i al shoot and s h ow no l a t e r al b e n d i n g. in an e v er s u d d e n ly b e g in a c t i ve g r o w th and f o rm s t i ll m o re r h i z o m e s. T he r e s u lt is t h at t he a r ea of s o il is p o p u l a t ed i n c r e a s i ng c i r c le of p l a n ts t h at c o n t i n u a l ly m o v es o u t w a r ds f r om t he p a r e n t. In t he m e a n t i m e, t he o l d er p l a n ts at t he c e n t er of t he c i r c le a re i n t e r- c o n t i n u a l ly p r o d u c i ng n ew r h i z o m e s. T h u s, t he s o il b e c o m es l a c ed w i th a d e n se m at of p l a n ts and r h i z o m es w h i ch b e c o m es c o n- s t a n t ly d e n s er at t he c e n t er u n t il t he c h a r a c t e r i s t ic b l u e g r a ss s od is f o r m e d. F ew p l a nt s p e c i es c an c o m p e te w i th t h is d o m i n e e r i ng g r o w th p a t t e r n. S u m m a ry T he r h i z o me of K e n t u c ky b l u e g r a ss is not t he lax, a i m l e s s ly g r o w i n g, u n d e r g r o u nd s t em of p o p u l ar m i s c o n c e p t i o n. h i g h ly o r g a n i z ed s t r u c t u r e, s u b j e ct to m a ny o r i e n t a t i o ns with a c h a n g i ng m o r p h o l o gy t h r o u g h o ut i ts v a r i o us g r o w th p h a s e s. It is a It a r i s es as a bud p r i m o r d i um in a p r e d e t e r m i n ed l o c a t i on d i c t a t ed by t he p a r e nt shoot a p i c al m e r i s t e m. It b e n ds d o w n w a r ds i n to t he s o il w h i le r o t a t i ng i ts p l a ne of s y m m e t ry by r e a s on of b i o- l o g i c al i n f l u e n c es d e r i v ed f r om i ts p a r e nt stfem. w i t h in t he s o il a re d e t e r m i n ed (a) v e r t i c a l l y, by g r a v i t a t i o n al s t i m- u l us on u n k n o wn b i o l o g i c al s e n s o r s, and (b) h o r i z o n t a l l y, by s t r i c t ly b i o l o g i c al c o n t r o l s. p l i c a t ed m o t i on in w h i ch one h e l i c al m o v e m e nt of s m a ll a m p l i t u d e, but r a p id r o t a t i o n, is s u p e r i m p o s ed on one of l a r ge m a g n i t u de but s l o w er r o t a t i o n. F i n a l l y, it t u r ns u p w a r ds t o w a r ds t he s o il s u r f a ce w i th i ts m o r p h o l o gy c h a n g i ng in p r e p a r a t i on f or i ts u l t i m a te as an a b o v e - g r o u nd s h o o t. It b o r es t h r o u gh t he s o il u s i ng a h i g h ly c o m- Its o r i e n t a t i o ns It is an e x c e e d i n g ly c o m p l ex o r g a n. r o le R e f e r e n c es C i t ed E t t e r, A. G. 1951. How K e n t u c ky b l u e g r a ss g r o w s. Ann. M i s s o u ri Bot. G a r d e n, .38: 2 9 3 - 3 7 5. E t t e r, A. G. 19 53. B l u e g r a ss p a s t u re a l m a n a c. Ann. M i s s o u ri Bot. G a r d e n, 40: 1 - 3 0. E v a n s, M. W. 1949. K e n t u c ky b l u e g r a s s. Ohio A g r. E x p t. Sta. R e s. Bull. 681. F i s h e r, J. E. 19 64. E v i d e n ce of c i r c u m n u t a t i o n al g r o w th m o v e- m e n ts of r h i z o m es of P oa p r a t e n s is L. t h at aid in s o il p e n e- t r a t i o n. Can. J. Bot. 42: ~293~-~299. F i s h e r, J. E. 19 65. M o r p h o l o g i c a l ly d i s t i n ct s t a g es in t he g r o w th and d e v e l o p m e nt of r h i z o m es of P oa p r a t e n s is L. and t h e ir c o r r e l a t i on w i th s p e c i f ic g e o t r o p ic r e s p o n s e s. Can. J. Bot. 43: 1 1 6 3 - 1 1 7 5. F i s h e r, J. E. 19 66a. T he g r o w th of r h i z o m es in K e n t u c ky b l u e- I n i t i a t i on and e a r ly g r o w t h. G r e e n h o u s e - G a r d e n, g r a ss I. G r a s s, 5(3): 1 - 7. F i s h e r, J. E. 1966b. T he g r o w th of r h i z o m es in K e n t u c ky b l u e- gra(ss II. F o rm and s t r u c t u re d u r i ng t he m a in s t a g es of g r o w t h. G r e e n h o u s e - G a r d e n, G r a ss 5(4): 1 - 6. F i s h e r, J. E. 19 72. S t r u c t u r al p o s i t i o n i ng and o r i e n t a t i o n al d e v e l o p m e nt in t he v e g e t a t i ve o r g a ns of P oa p r a t e n s is L. with s p e c i al r e f e r e n ce to t he r h i z o me a x i l l a ry b u d s. Can. J. Bot. 50: 7 4 3 - 7 5 0. L a r s o n, P. 19 62. G e o t r o p i s m, an i n t r o d u c t i o n. In H a n d b u ch d er P f l a n z e n p h y s i o l o g i e. Vol. 17, Sect. 2. W. R u h l a nd ( e d .) S p r i n g e r - V e r l a g, B e r l i n, pp. 3 4 - 7 3. S e r e b r y a k o v a, T. I. 19 68 B r a n c h i ng and T i l l e r i ng in t he P o a c e ae ( T r a n s l a t i on by f a m i l y. B o t a n i c h e s ky Z h u r n al _54: 8 5 8 - 8 7 1. t he F o r e i gn L a n g u a g es D i v i s i o n, D e p t. of t he S e c r e t a ry of S t a t e, G o v e r n m e nt of C a n a d a ). S h a r m a n, B. C. 1942. D e v e l o p m e n t al a n a t o my of t he shoot of Z ea m a ys L. Ann. Bot. ( London), N. S. 6: 2 4 5 - 2 8 2. S h a r m a n, B. C. Bot. Gaz. 1945. L e af and bud i n i t i a t i on in t he G r a m i n e a e. 106: 2 6 9 - 2 8 9. S h a r m a n, B. C. 1947. T he b i o l o gy and d e v e l o p m e n t al m o r p h o l o gy of t he shoot a p ex in t he G r a m i n e a e. N ew P h y t o l o g i s t, 2 0 - 3 4. 39: T U R F G R A SS R E S E A R CH IN B R I T I SH C O L U M B IA D o u g l as K. T a y l or R e s e a r ch S t a t i on C a n a da D e p a r t m e nt of A g r i c u l t u re A g a s s i z, B . C ., C a n a da I n t r o d u c t i on A t u r f g r a ss r e s e a r ch p r o g r am w as i n i t i a t ed at A g a s s i z, B. C., in 1969 by t he R e s e a r ch B r a n c h, C . D . A. T h is is a n ew p r o g r am f or t he A g a s s iz R e s e a r ch S t a t i on w h i ch is l o c a t ed 70 m i l es e a st of V a n- c o u v e r, B. C., at t he e a s t e rn end of t he L o w er F r a s er V a l l e y. A g a s s iz h as a m a r i t i me c l i m a te w i th an a n n u al p r e c i p i t a t i on of 64.3 in. w h i ch i n c l u d es 33 in. of snow. W i n t e r k i ll is r a r e, but w h en it r y e g r a s s. o c c u rs it is u s u a l ly c o n f i n ed to b e n t g r a ss and p e r e n n i al In a d d i t i on to t he A g a s s iz l o c a t i o n, c o o p e r a t i ve u n i f o rm t e s ts a re c o n d u c t ed at t he K a m l o o ps R e s e a r ch S t a t i on in t he i n t e r i or of t he p r o v i n ce w h e re c o n d i t i o ns a re c l o s er to a c o n t i n e n t al t y pe of c l i- m a t e. C o ol s e a s on g r a s s e s, f i ne f e s c u e s, b e n t g r a s s es and p e r e n n i al r y e g r a s s es a re w e ll a d a p t ed to C o a s t al B r i t i sh C o l u m b i a. A l t h o u gh t he K e n t u c ky b l u e g r a s s es a re not as p e r s i s t a n t, t h e re is n e ed f or t h is s p e c i es in h o me l a w ns a nd s p o r ts t u rf b e c a u se of its e a se of m a n a g e m e nt and i ts w e ar r e s i s t a nt q u a l i t i e s. H o w e v e r, in t he i n t e r i or of t he p r o v i n c e, K e n t u c ky b l u e g r a ss and f i ne f e s c u es a re b o th w e ll a d a p t e d. M a j or d i s e a s es a re leaf s p o t - m e l t i ng out c o m- p l ex ( H e l m i n t h o s p o r i um spp.), F u s a r i um p a t ch ( F u s a r i um n i v a l e ), s n o w m o ld ( T y p h u la spp. and F. n i v a l i) and r ed t h r e ad ( C o r t i c i um f u c i f o r m e ). L e ss f r e q u e nt a re O p h i o b o l us p a t ch ( O p h i o b o l us g r a- m i n is v a r. a v e n a e ), R h i z o c t o n ia b r o wn p a t ch ( R h i z o c t o n ia s o l a n i ), t h at f a i ry r i n g, leaf r u st and p o w d e ry m i l d e w. o ur s p e c i es and v a r i e ty p e r f o r m a n c es r e f l e ct r e s i s t a n ce to t h e se d i s e a s e s. It is q u i te e v i d e nt V a r i e ty T r i a ls V a r i e t al e v a l u a t i on h as b e en a m a j or p a rt of t he A g a s s iz t u r f- g r a ss p r o g r am to d a t e. S e e d i n gs m a de in 19 69 a nd 19 70 i n c l u de a t o t al of 242 e n t r i es in r e p l i c a t ed t r i a l s. F i ne f e s c u e s, K e n t u c ky b l u e g r a s s es and m i x t u r es a re e v a l u a t ed at t wo h e i g h ts of cut, 3 /4 in. and 1 l /2 in.; p e r e n n i al r y e g r a s s es at 1 l /2 in., and b e n t g r a s s es at 1/4 in.. F i ne f e s c ue A m o ng t he c r e e p i ng r ed g r o u p, D a w s o n, S59, R a s e n g o ld and L eo h a ve a p p r o a c h ed P e n n l a wn in p e r f o r m a n c e. S A I - 67 f r om S c o t l a nd is p a r t i c u l a r ly p r o m i s i ng in a p r e l i m i n a ry t r i a l. T he c h e w i n gs f e s c u e s, as a g r o u p, a re r e l a t i v e ly m o re a t t r a c t i ve with s u p e r i or d e n s i ty and r e s i s t a n ce in s o me c a s es to r ed t h r e a d. In p a r t i c u l a r, K o k e t, H i g h l i g h t, R o l ax and W i n t e r g r e en a re p r o m i s i n g. J a m e s t o wn is o u t s t a n d i ng f or i ts d a rk g r e en c o l or w h i le G o l f r o od s h o ws r e s i s t a n ce to F u s a r i um p a t c h. C - 2 6, a d a rk g r e en h a rd f e s- c u e, h as b e en s u p e r i or to a ll f e s c u es t e s t ed f or i ts d e n s i ty and r ed t h r e ad r e s i s t a n c e. K e n t u c ky b l u e g r a ss T he p e r f o r m a n ce of t h is s p e c i es h as b e en d i r e c t ly r e l a t ed to v a r i e t al r e s i s t a n ce to t he leaf s p o t - m e l t i ng out c o m p l e x. T he m e l t i ng out p h a se of t h is d i s e a se h as b e en so s e v e re t h at in p l o ts of c e r t a in v a r i e t i e s, w e e ds h a ve t a k en o v er 25% of t he plot w i t h in a t h r ee m o n th p e r i o d. M e r i on h as b e en m o st r e s i s t a nt to d a t e, f o l l o w ed by N u g g e t. O t h er v a r i e t i es t e s t ed h a v i ng b e t t er a v e r a ge r e s i s t a n ce a re A34, B a r o n, B i r k a, B101, E VB 391, F y l k i n g, Golf, K412, N e w p o r t, P e n n s t a r, Sodco, S y d s p o rt and W i n d s o r. T he i n c i d e n ce of H e l m i n t h o s p o r i um a p p e ar to be an i d e al s c r e e n i ng a r ea f or n ew r e s i s t a nt v a r i e t i e s. is so g r e at t h at it would t h an spp. B e n t g r a ss A m o ng t he c o l o n i al b e n t g r a s s e s, B r a b a n t ia ( E v a t e ), T r a c e n ta and B a r d ot a re p r o m i s i n g. P e n n c r o ss c r e e p i ng b e n t g r a ss and K i n g s t o wn v e l v et b e n t g r a ss a re both o u t s t a n d i ng in t h e ir c l a s s. P e r e n n i al r y e g r a ss T wo v a r i e t i es h a ve b e en p r o m i s i ng to d a t e: N o r l ea f or i ts w i n t er h a r d i n e s s, and M a n h a t t an f or i ts s u p e r i or d e n s i ty and a t- t r a c t i ve d a rk g r e en c o l o r. T h e re is a g r o w i ng i n t e r e st in t he u se of p e r e n n i al r y e g r a ss f or i n t e n s i v e ly u s ed p l a y i ng f i e l d s. To d a t e, o ur p e r e n n i al r y e g r a ss t r i a ls h a ve not b e en s u b j e ct to w e a r, but t h is f a c t or w i ll be an e s s e n t i al p a rt of f u t u re t r i a l s. T e st p l o ts h a ve b e en e s t a b l i s h ed in 19 72 of p e r e n n i al r y e g r a s s e s, K e n t u c ky b l u e g r a s s e s, t i m o t h i e s, and m i s c e l l a n e o us g r a s s e s, both in p u re s t a n ds and in m i x t u r e s. E v a l u a t i on of t h e ir p e r f o r m a n ce and c o n- t r i b u t i on of s o me of t he i m p r o v ed K e n t u c ky b l u e g r a ss c u l t i v a rs u s i ng a w e ar t r e a t m e nt d u r i ng t he p e r i od of m a x i m um u se is a m a j or goal. M i x t u r es T r i a ls a re u n d e r w ay a i m ed at p r o v i d i ng i n f o r m a t i on f or h o me l a wn r e c o m m e n d a t i o n s. U n d er t r i al a re P e n n l a wn and B o r e al f e s- c u e, M e r i on and P a rk K e n t u c ky b l u e g r a ss and H i g h l a nd b e n t g r a ss in a ll c o m b i n a t i o ns and at s e v e r al s e e d i ng r a t i o s. T he v i g or and c o m p e t i t i on f r om H i g h l a nd is m o st e v i d e nt and it a p p e a rs t h at t he c o n t r i b u t i on of t he f e s c ue or K e n t u c ky b l u e g r a ss c o m p o n e nt is d e- p e n d e nt u p on v a r i e t y. W i t h in t h r ee y e a r s, H i g h l a nd d o m i n a t es m o st m i x t u r e s. A l t h o u gh w e ll a d a p t e d, H i g h l a nd p r e s e n ts a m a n a g e m e nt p r o b l em to t he h o m e o w n e r, who a l m o st i n v a r i a b ly c u ts h is l a wn at a h e i g ht n o r m a l ly r e c o m m e n d ed f or K e n t u c ky b l u e g r a s s. F e r t i l i ty T r i a ls A c o m p a r i s on of e f f e c ts of s o u r c es and l e v e ls of n i t r o g en and s o me m i n or e l e m e n ts in. c u t t i ng h e i g ht is n ow in i ts t h i rd y e a r. A l e ss e x t e n s i ve t r i al is a l so u n d e r w ay to i n v e s t i g a te t he c u m u l a t i ve e f f e c ts of s e v e r al l e v e ls of p h o s p h o r o us and p o t a s s i um on S e a s i de b e n t g r a s s. in P e n n c r o ss b e n t g r a ss m a i n t a i n ed at l /4 H i gh Sand G r e e ns M i x es An i n s t a l l a t i on h as b e en m a de to o b s e r ve t he r e s p o n se of s ix v e g e t a t i ve b e n t g r a ss c u l t i v a rs g r o w i ng on s a n d - o r g a n ic m a t- t er m i x t u r e s. V a r i a b l es u n d er o b s e r v a t i on a re t wo d e p t hs of s a n d, s a w d u st v s. p e a t, and t h r ee d e p t hs of o r g a n ic m a t t er m i x- t u r e. We a re hoping to o b t a in i n f o r m a t i on on t he i r r i g a t i on r e- q u i r e m e n ts and r e l a te t h e se to e v a p o - t r a n s p i r a t i on and o t h er c l i m a t o l o g i c al d a t a. D i s e a se C o n t r ol E x p e r i m e n ts in s n ow m o ld ( m o s t ly T y p h u l a s p p .) c o n t r ol h a ve b e en i n i t i a t ed on t h r ee golf c o u r s es in t he i n t e r i or of t he p r o v i n c e. D e m o s an and P C NB a re t he m o st p r o m i s i ng t r e a t- m e n ts a f t er one y e ar of t r i a l s, but c e r t a in t r e a t m e nt x l o c a t i on i n t e r a c t i o ns a re a p p a r e n t. T U R F G R A SS I M P R O V E M E NT AT MICHIGAN S T A TE K e n y on T. P a y ne D e p a r t m e nt of C r op and Soil S c i e n c es M i c h i g an S t a te U n i v e r s i ty E a st L a n s i n g, M i c h i g an 48823 I n t r o d u c t i on O ur c l i m a te in M i c h i g an is p e r h a ps d i f f e r e nt t h an s o me of you m ay r e a l i ze in t h at we a re f r e q u e n t ly s u b j e ct to d r o u g h t. U s u a l ly e v e ry t h r ee out of f o ur y e a rs we c an count on q u i te a s e v e re d r o u g ht p e r i od d u r i ng t he s u m m e r. We get 30 to 35 in. of p r e c i p i t a t i on a n n u a l l y, w h i ch n o r m a l ly would s e em to be enough. D ry c o n d i t i o n s, h o w e v e r, a re t he r u le r a t h er t h an t he e x c e p t i on in m a ny s a n dy s o il a r e as d u r i ng m o st of o ur s u m m e r s. W i th o v er 25% of o ur l a w ns in s h a d ed s i t u a t i o n s, we n e ed a good t u r f g r a ss with both d r o u g ht and s h a de t o l e r a n c e. B a c k g r o u nd A n u m b er of p e o p le a re doing f i ne w o rk in b l u e g r a ss v a r i e ty t u r f g r a s s es is q u e en of t he d e v e l o p m e n t. T h is s p e c i e s, of c o u r s e, f or cool, h u m id r e g i o n s. W h en t he o p p o r t u n i ty c a me to e s t a b l i sh a b r e e d i ng p r o g r am t h r ee or f o ur y e a rs ago at M i c h i g an S t a te U n i v e r s i t y, it w as d e c i d ed t h at p e r h a ps t he f i ne f e s c u es h ad t he m o st to o f f er in p r o v i d i ng w h at is n e e d ed in t e r ms of s h a de and d r o u g ht t o l e r a n c e, as w e ll as t he a b i l i ty to g r ow with r e l a t i v e ly l o w er l e v e ls of f e r t i l i z er and m a n a g e m e n t. A c c o r d i n g l y, w h en t he d e c i s i on w as m a de to go i n to a b r e e d i ng p r o g r a m, it w as d e- c i d ed to i m p h a s i ze t he f e s c u e s. D r. F r ed E l l i o tt w as doing f o r- a ge b r e e d i ng w o rk and h ad i n c l u d ed s o me w o rk with t he f e s c u e s. As a r e s u l t, w h en t he p r e s e nt p r o j e ct w as i n i t i a t ed in 1968, a f i e ld of about 300 s i n g le p l a n ts of f i ne f e s c u es e x i s t ed t h at D r. E l l i o tt h ad a c q u i r ed t h r o u gh t he P l a nt I n t r o d u c t i o n. Of t h is g r o u p, t h e re w e re only t wo t h at w e re s t r o ng c r e e p e r s. One w i th t he w i de l e a f, a p p a r e n t ly 56 c h r o m o s o me c o m p l e x, and t he o t h er r e l a t i v e ly n a r r ow l e a f e d, c r e e p i n g - t y p e. F r om s u b s e q u e nt o b- s e r v a t i o n s, t he n a r r ow l e a f e d, c r e e p i n g - t y pe is not c o m m o n ly f o u n d. T h e re a re s o me v e ry a c c e p t a b le f e s c ue l a w ns in s h a d ed a r e as in m a ny of o ur h o me c o n d i t i o n s, but t h e re a re v e ry f ew e x c e l l e nt f e s c ue l a w ns in f u ll s u n l i g h t. L e af spot b e c o m es so s e v e re s e e d i ng a r ed f e s c ue l a wn is not r e c o m m e n d e d. S o m e t i me d u r i ng t he s u m m e r, it w i ll b e c o me i n f e c t e d. As a r e s u l t, it w as d e c i d ed to c o n c e n t r a te on leaf spot r e s i s t a n ce and s p r e a d i ng habit to p r o- v i de a v a r i e ty t h at s od g r o w e rs as w e ll as h o m e o w n e rs could u s e. M i c h i g an h as a l a r ge s od g r o w i ng i n d u s t r y. be a $ 3 5 - 40 m i l l i o n / y e ar c r o p. T he sod p r o d u c er n e e ds t h i ng t h at w i ll c r e ep r e l a t i v e ly q u i c k ly in o r d er to p r o v i de a m o re r a p id t u rn o v er t h an c an be o b t a i n ed f r om p r e s e nt f e s c ue v a r i e t i e s. is e s t i m a t ed s o m e- t h at r ed to It B r e e d i ng M a t e r i a ls and P r o c e d u r es U s i ng t he t wo p l a n ts f r om D r. E l l i o t t 's n u r s e ry as b a s ic t h r o u gh m a t e r i a l s, a c r o s s i ng n u r s e ry w as e s t a b l i s h ed and p l a nt a c c e s s i on and o t h er s o u r c e s, g e rm p l a sm w as a c c u m u l a- t e d. A y e ar l a t e r, D r. J o s e ph V a r g as f i n i s h ed h is P h . D. at t he U n i v e r s i ty of M i n n e s o ta and c a me to M i c h i g an S t a te as a f u l l - t i me t u r f g r a ss p a t h o l o g i s t. A s e t up w as t h us a v a i l a b le to b e g in s c r e e n i ng and a c t i v e ly w o r k i ng on t he d i s e a se p h a s es of t h is p a r t i c u l ar p r o j e c t. A f t er t wo or t h r ee g e n e r a t i o n s, ing h a b i ts i m p r o v ed c o n s i d e r a b l y. W h i le m o st of t he c r e e p i ng p l a n ts a re of t he w i d er leaf t y p e, we a re not as c o n c e r n ed about t h is as t he E u r o p e an g r a ss b r e e d e rs b e c a u se it is f e lt t h at w i de leaf w i ll b l e nd b e t t er w i th b l u e g r a s s es as t u r f. t he s p r e a d- A s c r e e n i ng p r o g r am w as e s t a b l i s h ed t h r ee y e a rs ago and D r. V a r g as h ad q u i te a bit of d i f f i c u l ty in t he i n i t i al p h a s es f i n d i ng t he r i g ht s p o re l o ad w i th w h i ch to i n o c u l a te t h e se f e s c u e s. With s p o re l o a ds w h i ch h ad b e en u s ed in M i n n e s o ta in t he b l u e- g r a s s e s, a ll p l a n ts b e c a me i n f e c t e d. T he n e xt s t ep w as to f i nd a c o n c e n t r a t i on of s p o r es s u f f i c i e nt to i s o l a te a f ew s u r v i v i ng f e s c ue p l a n t s. In a d d i t i on to c o m i ng down to a 30,000 s p o r e / cc in load, he f o u nd t h at if he s u s p e n d ed t he s p o r es in an a g ar s l u r r y, w h i ch w as t h en s p r a y ed on t he p l a n ts and p l a c ed in a m i st c h a m b er f or 48 h o u r s, v e ry e x c e l l e nt i n n o c u l a t i o ns would be o b t a i n ed and a f ew t o l e r a nt p l a n ts could be i s o l a t e d. With t h is t e c h n i q u e, f r om t he m a t e r i a ls we w e re s t a r t i ng with, h a v i ng no i n d i c a t i o ns or h i n ts as to w h e t h er t h e re w as a ny r e s i s t a n ce at a ll in t h e m, we i s o l a t ed about 1%. T h e se w e re i n o c u l a t ed a s e c o nd t i me and we k n o c k ed out a b o ut 90% of t h o s e. As a r e s u lt of t h r ee w i n t e rs of s c r e e n i n g, 18 c l o n es s u r v i v ed t h at w e re i n o c u l a t ed a t h i rd t i me a f t er t h ey had g r o wn in t he f i e ld t he w i n t er of 19 71-72. T wo c l o n es w e re f r om C - 2 6, w h i ch is t he m o st t o l e r a nt v a r i e ty and m i g ht be e x p e c t ed to p r o v i de c o n s i d e r a b le leaf spot r e s i s t a n c e. S u r p r i s i n g l y, it d i d n ' t. T h r ee or f o ur s u r v i v o rs a re f r om D r. R e ed F u n k 's c o l- l e c t i o ns -- F -3 m a t e r i al and m a t e r i al t h at h e 's c o l l e c t ed in t he E a s t. One is a c h e w i n gs t y p e, w h i ch is f r om an a c c e s s i on w h i ch a p p e a rs to be a n o t h er s p e c i e s, and t he r e st a re of t he b r o a d - l e a f ed c r e e p er t y p e s. T h e se a re b e i ng i n t e r c r o s s ed in t he g r e e n h o u se as w e ll as in t he f i e ld t h is s u m m er (19 72) and, by c o n t i n u ed s e- l e c t i on p r e s s u r e, be o b t a i n e d. t h at v e ry f ew h a ve s u r v i v ed t h is v e ry s e v e re t e s t, t h at w h i le we d o n 't h a ve t he r e s i s t a n ce t h at M e r i on b r o u g ht to t he b l u e g r a s s e s, s o me p r o g r e ss It is f e lt t h at w i th t h is s t r o ng s c r e e n i ng and t he f a ct it is h o p ed t h at a n ew p l a t e au is b e i ng m a d e. of r e s i s t a n ce c an A w i n t er h a r dy m e a d ow f e s c ue h as b e en d e v e l o p ed w h i ch h as s u r v i v ed v e ry w e ll in c o n t r a st to t he 50% s u r v i v al t h at p r e- v a i ls with K e n t u c ky 31, Alt a. a nd o t h er t a ll f e s c u es in M i c h i g an In a n o t h er p l a n t i ng m a de l a st f a ll in c o n j u n c t i on w i th M e r i on it h ad 100% s u r v i v a l. Alta t a ll f e s c ue K e n t u c ky b l u e g r a s s, went out 50% t h is w i n t er (19 71-72). T wo t h o u s a nd p o u n ds of s e ed w e re p r o d u c ed in O r e g on in 1971, and t h is is n ow in t r i a l. T h is could be u s e f ul on a l a r ge i n d u s t r i al l a wn or l a r ge s i te w h e re a v e ry f i ne h i g h ly g r o o m ed g r a ss but w h e re u n i f o r m i ty is d e s i r a b l e. of t he p a r e n t al m a t e r i a l, t h at it w i ll s u s t a in i t s e lf in c o m b i n a t i on w i th M e r i on b l u e g r a ss v e ry w e l l. T he leaf w i d th m a de it d i f f i c u lt to d i s t i n g u i sh f r om M e r i o n, a l t h o u gh it c an d u r i ng t he w i n t er w h en t he c o l or p a t t e r ns a re s o m e w h at d i f f e r e n t. It h a s n 't b e en t e s t ed f or w e ar i s n 't n e c e s s a r i ly n e e d e d, t o l e r a n c e. It a p p e a rs f r om t he p e r f o r m a n ce t e s t s. be d i s t i n g u i s h ed e a s i ly A s e r i es of t e s ts h as b e en c o n d u c t ed in a s h a de n u r s e ry d e- v e l o p ed in I9 60 u n d er n o n - i r r i g a t ed c o n d i t i o n s. T h is l a st s u m m er (19 71), w a t er w as p i p ed to t he a r ea and a n ew s et of eight or n i ne e x p e r i m e n ts a re u n d er way. T he s h a de is about 9 0 -9 5%. T he p r e s e nt t e s ts i n c l u de m i x t u r es and b l e n d s, and a c o m p a r i s on of v a r i e t i es u s i ng c h e m i c al d i s e a se c o n t r o l. D r. V a r g as is u s i ng s p r a ys in s u c c e s s i on as t he s e a s on p r o- g r e s s e s, and is s p r a y i ng half of e a ch plot to t ry to c o n t r ol d i s e a s es d u r i ng t he s u m m e r. S o me v e ry i n t e r e s t i ng d i f f e r e n c es a re e v i d e n t. O t h er S t u d i es D r. J a m es B e a rd and h is g r a d u a te s t u d e n ts h a ve d e v e l o p ed an e x t r a c t i on t e c h n i q ue t h at s e e ms to be w o r k i ng v e ry w e ll in t he e l e c t r o p h o r e t ic i d e n t i f i c a t i on of p r o t e in c o m p o n e n ts in g r a ss v a r i e t i e s. H o p e f u l l y, t h is w i ll l e ad to v a r i e t al i d e n t i f i c a t i o n. Good r e p r o d u c t a b i l i ty b e t w e en t he bent v a r i e t i es h as b e en o b- t a i n e d. P r o g r e ss l i s h ed t h at t e st p l a n ts m u st be g r o wn in t he g r o w th c h a m b er u n d er q u i te t he s a me c o n d i t i o n s. W h en t h is is d o n e, v e ry good b r e a k o u ts of d i f f e r e nt p r o t e in f r a c t i o ns c an be o b t a i n e d. is b e i ng m a de with P o a. It s e e ms w e ll e s t a b- R E S E A R CH TO I M P R O VE M E T H O DS OF T E S T I NG V A R I E T AL P U R I TY L e R oy W. N i t t l e r, H e ad D e p a r t m e nt of Seed I n v e s t i g a t i o ns N. Y. S t a te A g r i c u l t u r al E x p e r i m e nt S t a t i o n, C o r n e ll U n i v e r s i ty G e n e v a, N ew Y o rk I n t r o d u c t i on W h en I s t a r t ed w o rk at G e n e va n i n e t e en y e a rs ago we w e re t e s t i ng v a r i e t i es in t he f i e l d, and one a s s i g n m e nt I w as g i v en w as to i m p r o ve t e s t i ng m e t h o d s. O b v i o u s l y, f i e ld p l o ts h a ve s o me m a j or d i s a d v a n t a g e s. F i r s t, t h ey u s u a l ly t a ke f r om s ix m o n t hs to a y e ar to c o n d u c t. S e c o n d, no m a t t er w h en t he s e ed is r e c e i v ed it is n e c e s s a ry to wait u n t il s p r i ng to s t a rt a t e s t. I s t a r t ed with f o r a ge M o st of my w o rk h as d e a lt w i th g r o w i ng s e e d l i n gs in g r e e n- h o u s es and g r o w th c h a m b e rs and t r y i ng to f i nd w a ys s e e d l i n gs of o ne v a r i e ty d i f f er f r om t h o se of a n o t h e r. and g r a in c r o p s, t h en s e v en y e a rs ago we a d d ed T h i r t e en y e a rs ago o ur w o rk b e c a me p a rt of a N o r t h e a st R e g i o n al p r o j e ct on t e s t i ng f or v a r i e t al p u r i t y. P e n n s y l v a n i a, N ew Y o rk and t he USDA a re t he only a g e n c i es i n v o l v ed with it now. m o re s t a t es w e re i n v o l v ed w h en t he p r o j e ct s t a r t e d. At t h at it d e a lt m a i n ly w i th t e s t i ng a l f a l fa v a r i e t i es Guy M c K ee of T he P e n n s y l v a n ia S t a te U n i v e r s i ty h as done m u ch of h is w o rk on c h e m i c al t e s ts of s e e ds and s e e d l i n g s. T h u s, at G e n e va we h a ve c o n c e n t r a t ed o ur w o rk on s e e d l i ng g r o w t h. in f i e ld p l o t s. D r. t u r f g r a s s e s. S e v e r al t i me S e e d l i ng G r o w th R e s e a r ch I h a ve d i v i d ed my r e s e a r ch on s e e d l i ng g r o w th r o u g h ly into s ix c a t e g o r i e s. T he f i r st a p p r o a ch w as to g r ow s e e d l i n gs and in w h i ch o b s e r ve t h em c a r e f u l ly to d e t e ct a ny c h a r a c t e r i s t i cs t h o se of one v a r i e ty d i f f er f r om t h o se of a n o t h e r. T he s e c o nd w as u se of c o n t r o l l ed e n v i r o n m e n t al c o n d i t i o n s, m a i n ly t u r e, p h o t o p e r i o ds and light i n t e n s i t y. We m a n i p u l a t ed t h e se t e m p e r a- to f i nd c o n d i t i o ns t h at would i n d u ce s e e d l i n gs to e x p r e ss t h e ir g e n e t ic d i f f e r e n c es in s o me o b s e r v a b le c h a r a c t e r i s t i c. T h i r d, we u s ed c h e m i c a l s, e i t h er as f o l i ar s p r a ys or as s o il t r e a t m e n t s, to s ee t h o se if s e e d l i n gs of one v a r i e ty m i g ht r e s p o nd d i f f e r e n t ly f r om of a n o t h e r. F o u r t h, we d e f o l i a t ed s e e d l i n gs and o b s e r v ed t he r e- g r o w th to s ee if t h is would r e s u lt in o b s e r v a b le v a r i e t al d i f f e r- e n c e s. F i f t h, we g r ew s e e d l i n gs in s a nd and u s ed v a r i o us n u t r i e nt s o l u t i o ns to d e t e r m i ne if v a r i e t i es would r e s p o nd d i f f e r e n t ly to a l a ck of v a r i o us m i n e r al n u t r i e n ts or to d i f f e r e nt l e v e ls of t h e m. T he s i x th a p p r o a ch w as a c c e l e r a t i on of b l o o m i ng or h e a d i ng to m a ke t h e se plant p a r ts a v a i l a b le f or o b s e r v a t i on in a s h o rt T h is c an h a ve s o me r e al a d v a n t a g e s. F or e x a m p l e, about 10% of v e r n al a l f a l fa p l a n ts s h o u ld h a ve y e l l ow f l o w e r s. If we should t e st a s e ed s a m p le l a b e l ed V e r n al and f i nd no p l a n ts with y e l l ow f l o w e rs we would k n ow it w as m i s l a b e l e d. T h e se a p p r o a c h es c an n e v er be s e p a r a t ed c o m p l e t e ly but t h ey m a ke c o n v e n i e nt c a t e g o r i es f or t he p u r p o se of t h is t a l k. F or i n s t a n c e, s e e d l i n gs f or c h e m i c al a p p l i c a t i o ns we m u st c o n s i d er t he p e r a t u r e, g r o w n. T h is is a l so t r ue w h en a p e r s on is s t u d y i ng e f f e ct of m i n- e r al n u t r i t i o n. t e m- light i n t e n s i ty and p h o t o p e r i o ds u n d er w h i ch t h ey a re t i m e. in g r o w i ng O b s e r v a t i on of S e e d l i n gs By c a r e f ul o b s e r v a t i on of s e e d l i n gs d u r i ng t he f i r st t h r ee or f o ur w e e ks of g r o w t h, it is o f t en p o s s i b le to d e t e ct c h a r a c t e r i s t i cs in w h i ch v a r i e t i es d i f f e r. A l f a l fa v a r i e t i es w e re f o u nd to d i f f er in s e r r a t i on of t he f i r st l e a f. Oat v a r i e t i es d i f f er in l e n g th of i n t e r n o d e s. K e n t u c ky b l u e g r a ss v a r i e t i es d i f f er in leaf width, g r o w th h a b it and l e n g th of leaf s h e a t h s. C o n t r o l l ed E n v i r o n m e n ts In t he s e c o nd a p p r o a c h, one of t he f i r st t h i n gs we did w as to d e v e l op a t e st f or d i s t i n g u i s h i ng h a r dy f r om n o n - h a r dy a l f a l f a. W h en we g r ew p l a n ts w i th s h o rt p h o t o p e r i o d s, n o n - h a r dy v a r i e- t i es d e v e l o p ed s t e ms t w i ce as long as t h o se of h a r dy v a r i e t i e s. W h en we u s ed not only s h o rt p h o t o p e r i o d s, but a l so cold d a rk p e r i o d s, s t e ms of n o n - h a r dy p l a n ts w e re f i ve t i m es as long as t h o se of h a r dy p l a n t s. When K e n t u c ky b l u e g r a ss s e e d l i n gs w e re g r o wn w i th h i gh light i n t e n s i ty and long p h o t o p e r i o d s, u s e f ul v a r i e t al d i f f e r e n c es d e v e l o p e d. Of t he 25 v a r i e t i es we t e s t e d, s e e d l i n gs of 20 d e v e l o p ed r ed l o w er leaf s h e a t h s. S e e d l i n gs of t he o t h er f i ve w e re e n t i r e ly g r e e n. R h i z o me and t i l l er d e v e l o p- m e nt could a l so be o b s e r v ed on t he s a me p l a n ts and t h e se d i f f e r ed c o n s i d e r a b ly b e t w e en v a r i e t i e s. an a v e r a ge of 5.2 t i l l e rs p er p l a n t, c o m p a r ed to N e w p o rt w h i ch d e v e l o p ed only 1.3 p er p l a n t. On t he o t h er h a n d, N e w p o rt had t w i ce as m a ny r h i z o m es p er p l a nt as M e r i o n. In one s u ch t e s t, M e r i on d e v e l o p ed C h e m i c al U se In t he t h i rd a p p r o a ch we h a ve f o u nd s e v e r al c h e m i c a ls u s e f ul f or d i s t i n g u i s h i ng v a r i e t i e s. An i l l u s t r a t i on is u se of e t h e p h on on F e s t u ca r u b r a. T h is r e s u l t ed in d e v e l o p m e nt of two d i s t i n c t ly d i f f e r e nt t y p es of p l a n t s. T he f i r st r e s p o n d ed r e l a t i v e ly l i t t le to t he c h e m i c al and leaf b l a d es w e re long and n a r r o w. S t e ms of t he s e c o nd t y pe e l o n g a t ed and t he leaf b l a d es w e re s h o rt and w i d e. R u by and R a n i er r ed f e s c ue had n e a r ly 100% s h o r t - w i de p l a n t s. P e n n l a wn and I l l a h ee e a ch had about 60% leaf p l a n t s. Highlight and J a m e s t o wn c h e w i n gs f e s c ue h ad 100% l o n g - n a r r ow leaf p l a n t s. l o n g - n a r r ow leaf M a l e ic h y d r a z i d e, a p p l i ed to s o il in w h i ch p e r e n n i al r y e g r a ss s e e d l i n gs w e re g r o w i n g, w as u s e f ul in d i s t i n g u i s h i ng M a n h a t t an f r om o t h er v a r i e t i e s. T h is c h e m i c al c a u s ed p l a n ts to d i e, but m o st M a n h a t t an p l a n ts s t a y ed g r e en l o n g er t h an t h o se of o t h er v a r i e t i e s. C a p t an a p p l i ed to s o il a f t er t i m o t hy s e e ds w e re p l a n t ed r e s u l t ed in s t u n t i ng of s o me p l a n ts and n o r m al g r o w th in o t h e r s. V a r i e t i es d i f f e r ed in p e r c e n t a ge of s t u n t ed p l a n t s. D e f o l i a t i on We f o u nd t h at r e m o v al of t he p r i m a ry s t em of b i r d s f o ot t r e- f o il s e e d l i n gs had a p r o f o u nd e f f e ct on g r o w th h a b it of s e c o n d a ry s t e m s. E m p i re p l a n ts w e re d e c o m b e n t, but Viking p l a n ts w e re u p r i g h t. C l i p p i ng a l so b r i n gs out g r o w th h a b it d i f f e r e n c es a m o ng a l f a l fa v a r i e t i e s. G r o w th h a b it of p e r e n n i al r y e g r a ss c an a l so be m o d i f i ed by clipping. N u t r i e nt M a n i p u l a t i on We s t a r t ed g r o w i ng s e e d l i n gs with n u t r i e nt s o l u t i o ns about In a d d i t i o n, a n u t r i e nt m ay be w i t h h e ld s ix y e a rs ago. T h is h as r e s u l t ed in m a ny i n t e r e s t i ng d i s c o v e r i e s. It is p o s s i b le to w i t h h o ld any p a r t i c u l ar n u t r i e nt a nd o b s e r ve f or d e f i c i e n cy s y m p t o m s. e i t h er b e f o re or a f t er a c o m p l e te n u t r i e nt s o l u t i on is u s e d. U se of d i f f e r e nt f o r ms and l e v e ls of n u t r i e n ts o f f e rs a d d i t i o n al p o s s i- b i l i t i e s. We p u b l i s h ed a p a p er r e c e n t ly on u«se of a s o l u t i on l a c k- ing c a l c i um w i th K e n t u c ky b l u e g r a ss v a r i e t i e s. S e e d l i n gs of s o me v a r i e t i es t u rn y e l l ow and d ie w h en t h ey a re g r o wn with t h is s o l u- t i o n. O t h e rs r e m a in g r e en a nd r e l a t i v e ly h e a l t h y. D e l ta and N u g g et a re v a r i e t i es t h at t u rn y e l l o w. A r i s t a, F y l k i ng and P e n n- s t ar r e m a in g r e e n. If M e r i on and W i n d s or a re g r o wn w i th a c o m p l e te s o l u t i o n, s e e d l i n gs of both v a r i e t i es h a ve g r e en leaf s h e a t hs and it is d i f f i- cult to s ee m u ch d i f f e r e n ce b e t w e en t h e m. H o w e v e r, if t h ey a re g r o wn with a c o m p l e te s o l u t i on t he f i r st t wo w e e ks and t h en with a s o l u t i on l a c k i ng n i t r o g en t he n e xt t h r ee w e e k s, l o w er leaf s h e a t hs of M e r i on b e c o me r ed and t h o se of W i n d s or r e m a in g r e e n. A s o- l u t i on l a c k i ng c a l c i u m, but with a high l e v el of n i t r o g en s u p p l i ed as a m m o n i um n i t r a t e, c a u s ed r ed c o l or to e x t e nd two i n c h es or m o re up Nugget p l a n t s. o n ly In t he s a me e x p e r i m e n t, r ed c o l or e x t e n d ed l /2 i n ch up B a r on p l a n t s. A c c e l e r a t i on of H e a d i ng or F l o w e r i ng By u se of c o n t i n u o us light and s u i t a b le t e m p e r a t u r e s, we c an to get a l f a l f a, b i r d s f o ot t r e f o i l, r ed c l o v er and s w e et c l o v er p l a n ts b l o om in 4 to 5 w e e ks f r om t he t i me t he s e ed is p l a n t e d. S p r i ng b a r l ey and o a ts w i ll h e ad out in 4 to 5 w e e ks u n d er s i m i l ar c o n- d i t i o n s. a n n u al f r om p e r e n n i al r y e g r a s s. B e c a u se t he f l u o r e s c e n ce t h at h as b e en u s ed in t he p a st is not a l w a ys r e l i a b l e, t h e re is n e ed f or a n ew m e t h od of d i s t i n g u i s h i ng t h e se s p e c i e s. We g r ew s e e d- l i n gs w i th c o n t i n u o us light, a c o m p l e te n u t r i e nt s o l u t i o n, and a In l a wn g r a s s e s, we h a ve u s ed t h is t e c h n i q ue to d i s t i n g u i sh t e st t e m p e r a t u re of 27°C f or 20 h o u rs a nd 16°C f or 4 h o u r s. W i t h in t h r ee w e e ks of t he t i me s e e ds w e re p l a n t e d, 10% of a n n u al p l a n ts h ad h e a d ed out and m o st of t he r e st h ad e l o n g a t ed s e ed s t a l k s. In c o n t r a s t, no p e r e n n i al r y e g r a ss p l a n ts had d e v e l o p ed a ny s t e ms at all. S u m m a ry In s p e c i es s u ch as K e n t u c ky b l u e g r a s s, b a r l ey and o a t s, v a r i e t i es t e nd to be v e ry u n i f o rm a nd in m a ny c a s es it is p o s s i b le to d i s t i n g u i sh o f f - t y pe p l a n ts w i th a high d e g r ee of p r e c i s i o n. In m o st c r o s s - p o l l i n a t ed s p e c i e s, p l a n ts d i f f er c o n s i d e r a b ly w i t h in a v a r i e t y. T h u s, it is d i f f i c u lt to d e v e l op r e l i a b le t e s ts f or d i s- t i n g u i s h i ng v a r i e t i es and it m ay be n e c e s s a ry to a c c e pt a l o w er d e g r ee of p r e c i s i o n. In a d d i t i on it m ay not be p o s s i b le to d i s t i n- g u i sh v a r i e t i es w i th s i m i l ar c h a r a c t e r i s t i c s. D i s c u s s i on P e r i od D r. D a n i e l: How long do you let y o ur s e e d l i n gs go in y o ur g r o w th h a b it t e s ts ? D r. N i t t l e r: In b l u e g r a s s, we h o pe to t e r m i n a te an e x p e r i m e nt f i ve w e e ks f r om t he t i me we s t a r t. We c an get good r e s u l ts on s o me of o ur oat t e s ts in t en d a ys by u s i ng a m i n u s - p h o s p h o r us s o l u t i o n. T h is r e s u l ts of s o me v a r i e t i es and not in o t h e r s. in d e v e l o p m e nt of r ed c o l or in leaf s h e a t hs in D r. P a y n e: We c e r t a i n ly would l i ke f or you to k e ep p u r s u i ng t h is m a t t er of f o r c i ng g r e e n h o u se w i n t er h e a d i ng in t he f e s c u e s. We t r i ed 16, 20, 2 4 - h o ur d ay r e g u l ar g r e e n h o u se c o n d i t i o ns a f t er a cold i n d u c t i o n, but d i d n!t o b t a in h e a d i n g. D r. N i t t l e r: T i m o t hy and s o me b e n t g r a s s es w i ll h e ad out in g r o w th c h a m b e r s, but m o st o t h er p e r e n n i al g r a s s es s u ch as r ed f e s c u e, o r c h a r d g r a s s, and p e r e n n i al r y e g r a ss a p p a r e n t ly n e ed to be v e r- n a l i z e d. T h e se w i ll not e v en p r o d u ce s t e m s. E t h e p h on w i ll r ed f e s c ue s e e d l i n gs to p r o d u ce s t e m s. T he f i r st t i me I s aw t h is I t h o u g ht t h ey would a l so h e ad out, but t h ey d i d n ' t. It m ay be t h at u se of e t h e p h on and cold d a rk p e r i o ds would i n d u ce h e a d i n g. T h is is s o m e t h i ng we p l an to i n d u ce t r y. D r. D u i c h: H a ve you e v er t r i ed a h e a vy m e t al a p p r o a c h? D r. N i t t l e r: No. D r. D u i c h: S o m e t i m es w i th f u n g i c i de a p p l i c a t i o n s, I k n ow q u i te a d i f f e r e n t i al c a d m i um r e s p o n se of v a r i e t i e s. T he P MA e f f e ct on M e r i on m i g ht be a good s y s t e m a t ic a p p r o a ch to t r y. t h e r e 's D r. N i t t l e r: T h e re a re so m a ny a g r i c u l t u r al c h e m i c a ls t h at is no l i m it to h ow m u ch w o rk you c an do w i th t h e m. T he g r o w th r e g u l a t o r s, out v a r i e t al d i f f e r e n c e s. in g e n e r a l, h a ve b e en a u s e f ul g r o up f or b r i n g i ng t h e re In T e x as a f ew y e a rs a go we n o t ed t h at c e r t a in D r. Long: c i d es would r e a ct d i f f e r e n t ly on s o me of t he b e r m u d a s. One a c t i on w as c h l o r d a ne i n t e r a c t i on on T e x t u rf 10 b e r m u d a. t u rn it v e ry yellow. i n s e c t i- i n t e r- It would D r. N i t t l e r: T h is is s o m e t h i ng I h a v e n!t l o o k ed at. S o me of t he f u n g i c i d e s, s u ch as c a p t a n, would be u s e f ul on c e r t a in g r a s s e s. G e n e r al D i s c u s s i on D r. Long: D r. F i s h e r, you m e n t i o n ed t h at if you h a ve a l l o w ed m o re p h o t o s y n t h e t ic a r ea on b l u e g r a s s es t h at you would p r o- b a b ly e n c o u r a ge r h i z o m i n g. D r. D a n i el s t a t ed t h at he h ad s p a ce p l a n ts t h at w e r e n 't c l i p p ed down, and got p r e t ty good r h i z o m i n g. Do you t h i nk t h at if t h e se w e re u n d er a l i t t le m o w i ng p r e s s u r e, t h ey w o u l d n 't r h i z o me as m u c h? D r. F i s h e r: T h a t 's o ur e x p e r i e n ce and a l so t he e x p e r i e n ce of a n u m b er of o t h er p e o p le who h a ve w o r k ed with r h i z o m e s, e v en of d i f f e r e nt s p e c i e s. If you cut t he p h o t o s y n t h e t ic a r ea of y o ur p l a nt down, r h i z o me p r o d u c t i on and r h i z o me g r o w th d r o ps off r a p i d l y. T he r h i z o me is a p a r a s i te on t he p a r e nt p l a nt a nd d e- p e n ds on an e x c e ss of p h o t o s y n t h a t e. e v i d e n ce t h at h i gh c a r b o h y d r a te l e v e ls w i th low n i t r o g en is t he b e st c o m b i n a t i on f or i n d u c i ng r h i z o me p r o d u c t i on c o u c h g r a s s. M a c l n t y re h as l e v e ls in In f a c t, D r. P a y n e: D r. D u i c h, h ow a re you h a n d l i ng t he b r e e d e r 's of P e nn S t a te v a r i e t i es as t h ey a re s e nt W e st f or i n c r e a s e? e x a m p l e, P e n n c r o ss m a i n t a i n i ng t h e se c l o n e s? s e ed F or is c o m p o s ed of t h r ee c l o n e s. How a re you D r. D u i c h: W e ' re m a i n t a i n i ng s p a ce p l a nt n u r s e r i es — a l t e r n a te r o w s. D r. P a y n e: How l a r ge a n u r s e ry do you m a i n t a in of t h o se c l o n e s? D r. D u i c h: s e ed n e e d s. About t wo a c r e s, d e p e n d i ng on p r o j e c t ed f o u n d a t i on D r. N i t t l e r: t he n e xt in P e n n f i n e, or w i ll it be p r e t ty s t a b l e? Is t h e re going to be a s h i ft f r om one g e n e r a t i on to D r. Duich: We h a v e n 't s e en a s h i ft in t wo g e n e r a t i o ns of s p a ce p l a nt and t u rf t e s t i n g. We c a n 't d i s t i n g u i sh a n y t h i n g. T h a t 's as f ar as w e ' ve gone. I m i g ht c o m m e n t, t o o, t h at w e ' re looking at t h is D r. N i t t l e r: in t he g r o w th c h a m b e rs t r y i ng to s ee if we c an d e t e r m i ne s o me d i f- f e r e n ce in t he r e l a t i v e ly young s t a g e. We t h i nk we c an s ee s o m e, e s p e c i a l ly in p l a nt s i z e. W e ' ve m e a s u r ed t he s e c o nd leaf b l a de l e n g t h. P e n n f i ne s e e ms to be s h o r t er t h an t he o t h er v a r i e t i es t h at you h a ve s h o w ed me in y o ur t a b l e s. A l s o, it t e n ds to h a ve l e ss r ed c o l or on t he l o w er leaf s h e a th t h an s o me of t he l a r g er v a r i e t i e s. D r. D a n i e l: D r. Long, h ow m u ch d w a r f n e ss do you t h i nk p e o p le w a nt in vt u r f g r a s s e s? D r. Long: a nd c l i p p i n g. To s t a rt with, a b o ut 70% r e d u c t i on in l a wn m o w i ng M r. M a y e r: How a re we going to m e a s u re d w a r f n e ss in a m o w ed l a w n? W e ' re doing it in a n u r s e r y, but d o es t h is c o r r e l a te with t he l a w n? T he h o m e o w n er m o ws h is l a wn at t wo i n c h e s. E v en if a m a t u re p l a nt g r o ws s ix i n c h e s, he is s t i ll going to m ow it at t wo i n c h e s. He is s t i ll going to be t a k i ng y i e l ds off. D r. D u i c h: t h o u g h, I t h i nk y o u ' ve a c h e i v ed s u c c e s s. If one h as to m ow it only t h r ee t i m es in t he s p r i n g, D r. D a n i e l: We s t a r t ed m e a s u r i ng y i e ld a f t er leaf spot h ad hit t he o t h er g r a s s e s. O ur h e a l t h i er g r a s s es p r o d u c ed t he m o st c l i p- p i n g s. T h o se t h at h ad b e en w e a k e n ed by t he d i s e a se p r o d u c ed l e ss c l i p p i n g s. T h is w as a c o m p l e t e ly e r r o n e o us p e r i od of o b s e r- v a t i o n. L e t 's a s s u me you get s o m e t h i ng down to w h e re y o u ' re m o w i ng t he g r a ss o n ce e v e ry two w e e ks at t h r ee q u a r t e rs of an inch, and y o ur c l i p p i ng l e n g th d o e s n 't e x c e ed y o ur m o w i ng h e i g h t. A re you c o m- m i t t ed t h en to k e ep out c r a b g r a ss or g r a s s es s u ch as a n n u al b l u e- g r a s s, r y e g r a ss and r ed f e s c ue w h en t h ey a re c o n s i d e r ed as w e e d s? Is t he l a n d s c a p er r e a dy to give t he h o m e o w n er t h at c l e an of a s e ed b e d? A re t he s e e d s m en r e a dy to c l e an up h is s e ed f i e ld w e ll enough, or is t he s e e d h o u se going to k e ep it c l e an so t h e re is no o t h er c r op in it so t h at it w i ll s t a nd to c o m p e t e? l o w - g r o w i ng t y p e s, t h at I 'd l i ke to m a ke one a s s o c i a t ed c o m m e nt f r om a I t h i nk w e ' ve b e en a l i t t le l ax in not D r. D u i c h: p r a c t i c al point of view. g e t t i ng s o me of t h is d a t a. W e ' ve n o t i c e d, p a r t i c u l a r ly on s o me of t h e se s o - c a l l ed d e c u m b e n t, s e e ms to be q u i te a v a r i e t al d i f f e r e n t i al a f t er you m ow on u n c o v e r ed f r om p r e v i o us m o w i n g s. S o me of t he r e m a i n i ng t u rn y e l l ow and b r o wn off. A p a r t i c u l ar v a r i e ty m ay look v e ry p o or u n t il you get e n o u gh r e g r o w th to c o v er it up. As s o on as you r e m ow it, e v en f r om t he p r e v i o us m o w i n g, you f i nd it a g a i n. t h i nk M e r i on h as b e en r a t h er n o t o r i o us f or t h is u n c l o se m o w i n g, h i g h - m a i n t e n a n ce v a r i e t i e s, N u g g e t, f or e x a m p l e, I d o n 't k n ow w h at h a p p e ns t he c l i p p i n g s. T h ey j u st t ic point of view, t r ue d w a r f n e s s, t h is could be a v e ry i m p o r t a nt c o n s i d e r a t i o n, t o o. l e v e l s. H o w e v e r, w i th s o me of t h e se o t h er to s e em to d i s a p p e a r. So f r om t he a e s t h e- If we would a c h e i ve t h ey look v e ry p r o m i s i n g. I t h e re l e a v es l e a v es D r. K u c: W h en you t a lk about a d w a r f, do you r e f er to t he u l t i- m a te h e i g ht of t he p l a n t, or do you r e f er to t he r a te of g r o w t h, or b o t h? Do a ll d w a rf p l a n ts h a ve a r e d u c ed r a te of g r o w t h? D r. D a n i e l: My i d ea of a d w a rf is one t h at h as 50% f e w er c l i p- If M e r i o n, p i n gs c o m p a r ed to what h as b e en n o r m al in t he p a s t. a s t a n d a r d, t h en a d w a rf v a r i e ty s h o u ld be about 50% l e s s. is giving you a c e r t a in y i e ld of c l i p p i n g s, D r. Long: D r. D u d e c k, w h at is y o ur e x p e r i e n ce w i th T i f d w a rf on a t t e m p ts to u se it in h o me l a wn s i t u a t i o ns in t he S o u t h e a s t? What a re t he p r o b l e ms w i th t h is d w a rf v a r i e t y? D r. D u d e c k: T i f d w a rf h as not b e en u s ed in t he h o me l a wn s i t u a- t i o n, to my k n o w l e d g e, b e c a u se of t he h i gh m a i n t e n a n ce i n v o l v e d. F l o r i da No Mow, FB 137 or F l o r a t u rf h as b e en p r o m o t ed and u s ed i n c o r r e c t l y, as t he n a me i m p l i e s, as a no m ow v a r i e t y. U n f o r- t u n a t e l y, t oo m a ny p r o b l e ms w e re f o u nd with t h is g r a ss in t he f i e ld p r i or to i ts r e l e a s e. A c t u a l l y, it h as n e v er b e en o f f i c i a l ly r e l e a s ed f r om F l o r i d a. T i f d w a r f, w h i ch d o es h a ve a s l o w er g r o w th r a te c o m p a r ed to t he o t h er b e r m u d a s, h o me l a wn s i t u a t i on b e c a u se of t he m a i n t e n a n ce is not u s ed in t he r e q u i r e d. In A r i z o n a, T i f d w a rf w i ll s e l d om get a b o ve t h r ee T i f d w a rf h as b e en u s ed a l i t t le in t he P h o e n ix a r ea D r. K n e e b o n e: as a s u b s t i t u te f or T i f g r e en with t he s a me i d ea as t he no m o w i ng s i t u a t i o n. w h e r e as T i f g r e en l e ft u n m o w ed m i g ht get up to eight i n c h e s. T i f- d w a rf r o l ls and s w i r l s. To be h a n d l ed p r o p e r ly it would be m o w ed at o n e - h a lf inch, w h i ch m e a ns m o w i ng j u st as f r e q u e n t ly as a ny o t h er b e r m u d a g r a s s. T h e re s h o u ld be e n o u gh g r o w th to self h e al a nd k e ep t he g r e en down so t he g r e en is b e l ow t he cut of any d w a r f. i n c h e s, B e r m u d a g r a ss d w a r fs a re d i f f e r e nt t h an t he b l u e g r a ss o n e s. m o re t he l e n g th of t he i n t e r n o de and l e n g th of t he i n d i v i d u al r a t h er t h an t he o v e r - a ll d w a r f n e ss of t he p l a n t. We get s o me a re m u ch m o re d w a rf t h an T i f d w a r f - - so d w a r fy t h at you c a n 't g r ow a l e a v es t h at lawn. I t 's I d o n 't t h i nk we g a ve D r. K uc a v e ry good a n s w er a while D r. L o n g: a go on w h e t h er w e ' re d e a l i ng w i th g r o w th r a t e, or g r o w th height r e s p o n s e. S o me y e a rs a go w h en I h ad a p r o g r am going and w as looking at s o me St. A u g u s t i ne d w a r f s, we did p i ck up t h is a s p e ct of g r o w th r a t e. C o m p a r i ng g r o w th r a te w i th one t á is w i th F l o r a t i ne or B i t t er B l ue o v er a p e r i od of f o ur m o n t h s, we m e a s u r ed no s t o l en g r o w th on t he d w a r f, as c o m p a r ed to 10 to 12 i n c h es of s t o l en g r o w th on F l o r a t i ne or B i t t er Blue. A p p a r- e n t ly we h a ve an e f f e ct on one a s p e ct of t he d w a r f n e s s. M a y be a s l o w i ng up of t he g r o w th r a t e. One of D r. D a n i e l 's s t u d e n ts f o u nd t h is in s o me of h is w o rk w i th b l u e g r a s s. He h ad v e ry g r o w th on s o me of t he m a t e r i al he w as s t u d y i n g. of t he e x p e r i m e n- l i t t le D r. D a n i e l: On t he o t h er end of t he s c a l e, one of t he f i r st p l a n ts we w o r k ed with, t he g r a n d m o t h er of 9 0% of o ur r e s e a r ch w o r k, w e ' ve n e v er f o u nd a n y t h i ng m o re h e t e r o g e n o us v i g o r o us in i ts own a b i l i ty to s p r e a d, t h an t he old o r i g i n a l - - t h is 16B we p i c k ed up off C h i c a go Golf Club. h a v e n 't f o u nd a n y t h i ng m o re v i g o r o us t h an t he o r i g i n al p l a n t, s e e m s, on s p r e a d. W e ' ve f o u nd a lot of d w a r f n e ss o v er t h is in i ts o f f s p r i n g, n or m o re In s e v en g e n e r a t i o n s, we it t i m e. D r. P a y n e: w a n t, we c an u se d w a r f n e ss if we c an get two or t h r ee v a r i e t i es In r e g a rd to y o ur q u e s t i on of h ow m u ch d w a r f n e ss we so t h at we c an get a b l e n d. But as you c an i m a g i n e, it p r o b a b ly w o n 't b l e nd t oo w e ll w i th o t h er t y p e s, e s p e c i a l ly w h en m o w i ng p a t t e r ns a r e n 't c o n s i s t a n t. we s h o u ld h a ve If we h a ve one, it would a p p e ar t h r e e. t h at J u st a c o m m e nt on t he p e r f o r m a n ce of Nugget, I ' ve n o t i c ed t h at in f a ll D r. D. T a y l o r: o ne of t he d w a rf K e n t u c ky b l u e g r a s s e s. p l a n t i n g, I ' ve h ad t r o u b le g e t t i ng a good s t a n d. With v a r i e t i es of t h is t y p e, if t he p r a c t i ce in t he a r ea is to f a ll p l a n t, w e ' ll h a ve to m o ve t he p l a n t i ng d a te a h e ad in o r d er to get a good e s t a b l i s h- m e nt b e f o re w i n t e r. l o n g er you w o u l d n 't h a ve to w o r ry as m u ch about I w o n d er if t h e re is s o me w ay to e d u c a te t he p u b l ic It would s e em to me t h at if t he p e o p le would cut t he D r. H i g g i n s: t he g r a ss d w a rf p l a n t s. to want and cut it t a l l e r, it would s i m p l i fy t he b r e e d i ng p r o b l em if you d i d n 't h a ve to w o r ry about d w a r f n e s s. T he f i r st q u e s t i on t h at c o m es to my m i nd is why do p e o p le want to cut it s h o r t? Is t h is b e c a u se t h ey h a te t h e ir l a wn a nd t h e y ' re t r y i ng to hold b a ck i ts g r o w th by d a m a g i ng t he g r a s s? Of c o u r se i t 's p h y s i o l o g i- I w o n d er if it m i g ht be e a s i er to s o l ve t he p r o b l em of c a l ly d o n e. m o re A m e r i c a ns h a v i ng b e t t er l a w ns if t h ey could be e d u c a t ed to r a i se t he b l a de a l i t t le bit, r a t h er t h an t ry to c o me up with v a r- i e t i es t h at a re m o re d w a r f. T h e r e 's e n o u gh p r o b l e ms of b r e e d i ng w i t h o ut h a v i ng to w o r ry about t h a t. D r. D a n i e l: As w e ' ve w o r k ed w i th o ur p l o t s, if we h a ve two h e i g h ts of cut, we n o r m a l ly s p e nd 9 5% of o ur t i me n o t i c i ng w h at h a p p e ns to t he low cut. If you want to u se t he t u rf f or c r o q u e t, f or a p a r t y, f or w a l k i ng or f or v i e w i n g, g e n e r a l ly t he m o re you c an m a n i c u re it, t he b e t t er it m ay p e r f o r m, p r o v i d i ng you h a ve s u r v i v a l. So if we c an d w a rf t he p l a nt and cut it at t h r ee q u a r t e rs of an i n c h, and cut off t h r ee q u a r t e rs of an inch, t h at m ay be as e q u a l ly s a t i s f a c t o ry to a l a wn as h a v i ng it two i n c h es high, and w h en m o w e d, c u t t i ng off t wo i n c h es of c l i p p i n g s. I n s t i t u t i o n al t u rf m a n a g e rs f r e q u e n t ly m a ke t he s t a t e- M r. B a n g s: m e nt t h a t, "I cut my t u rf as s h o rt as I c an so I c an r e d u ce n u m b er of t i m es t h at I h a ve to m ow i t ." if we could h a ve a c o m m e nt on t h at s t a t e m e nt b e c a u se it is q u i te c o m m o n. I t h i nk it m i g ht be h e l p f ul t he r e f er D r. F i s h e r: T he f o l l o w i ng s u g g e s t i on d o e s n 't n e c e s s a r i ly to a t u r f g r a ss c l o ne of b l u e g r a s s, but to t he v i g o r o us wild one t h at w e ' re w o r k i ng with. We f o u nd t h at w h en c u t t i ng f r e q u e n t ly and l o w e r, we had m u ch r e d u c ed g r o w t h. A f t er a p e r i o d, t he d ry w e i g ht a c c u m u l a t i on d r o p p e d, and we put it down to r e d u c i ng t he photo s y n t h e t ic t i s s ue q u i te d r a s t i c a l l y. d o es cut h is l a wn v e ry s h o rt d o e s n 't n e ed to cut it as o f t e n. He d o e s n 't h a ve to cut as m u ch m a t e r i a l. On t he o t h er hand, m a y be he d o es cut it as o f t en f or a p p e a r a n ce s a k e. T h e r e 's a big d i s- t i n c t i on h e re b e c a u se if he h as it t h at s h o r t, he m ay want to c o n- t i n ue m a i n t a i n i ng it s h o rt a nd would h a ve to m a n i c u re m o re o f t e n. But h e 's not cuting as m u ch m a t e r i al as t he m an who l e ts h is l a wn g r ow t h r ee i n c h es high. A f t er all, a h a l f - i n ch g r o w th d o e s n 't s h ow as m u ch on t h r ee i n c h es as it d o es on a half I t h i nk t he p e r s on t h at inch. D r. D a n i e l: Of c o u r s e, with a l a wn m o w er w e ' re j u st s i m u l a t i ng w h at t h e y ' ve done w i th b o n zi t r e es j u st a b o ut a ny s p e c i es and by t r i m m i ng both t op and r o o t, s u b j e ct it to g r o w th c o n t r o l. With a v a r i e ty of g r a s s, s h o rt cut is m o re d r o u g ht s u s c e p t i b le t h an a long cut on a g i v en day. So I 'm s u re we b o n zi b o th t op and r o ot as we t r im it down to a c o n s i d e r a b le extent. in J a p a n. T h ey c an t a ke I h a ve a c o m m e nt f r om a r a n ge s i t u a t i on D r. W h a l l e y: e x a c t ly a g r e es with w h a t 's b e en s a i d. c l ip r a n ge g r a s s e s, t h en t he r a te of b o th t op and r o ot g r o w th is c o n s i d e r a b ly l e s s e n e d. With a r e d u c ed r o ot s y s t e m, d r o u g ht r e s i s t a n ce is t h en d e c r e a s e d. In f a c t, I ' ve h e a rd it s a id with a l f a l fa t h at if you p u ll a leaf off t he top, a r o ot f a l ls off t he b o t t o m. I 'm s u re t h is is an e x a g g e r a t i o n, but t he p r i n c i p le c e r t a i n ly h o l d s. If you o v e r g r a ze or o v e r- t h at I ' ll s ay it f r om t he o p p o s i te point of view. W h en D r. M a d i s o n: we m o w ed a h e a l t hy t u rf at 2 l /2 i n c h es at i n t e r v a ls of two to t h r ee w e e k s, g r a ss v i g or built up so e x t e n s i v e ly t h at w i t h in two or t h r ee d a ys a f t er m o w i n g, an i n ch or t wo of g r o w th r e c o v e r ed a nd t he g r a ss a p p e a r ed r e a dy f or a n o t h er m o w i n g. BASICS IN GRASS I M P R O V E M E NT J o hn H. M a d i s on D e p a r t m e nt of E n v i r o n m e n t al H o r t i c u l t u re U n i v e r s i ty of C a l i f o r n ia D a v i s, C a l i f o r n ia 9 5616 I n t r o d u c t i on O f t en l a c k i ng in t u rf b r e e d i ng p r o g r a ms is t he bad e x a m p le a g a i n st w h i ch to c o m p a re t he m a ny f i ne s e l e c t i on and b r e e d i ng s t u d i es so t h at t h e ir e x c e l l e n ce c an be a p p r e c i a t e d. l i ke to give an e x a m p le of a p o or p r o g r am w h i ch a r o se u n i n t e n- t i o n a l l y. I s h o u ld if In 19 59, we a b a n d o n ed p l o ts of c r e e p i ng b e n t g r a ss ( A g r o s t is s t o l o n i f e r a) b e c a u se of c r o ss c o n t a m i n a t i on w i th b l u e g r a s s. We s p r a y ed t he p l o ts w i th w e ed o il and w i t h h e ld i r r i g a t i on w a t er f or t he s u m m e r. With f a ll r a i n s, m a ny g r a s s es r e c o v e r ed and g r e w. T he next s u m m er we not only w i t h h e ld w a t er but u s ed w e ed o il t w i c e, b u r n i ng t he f i e ld a f t er t he s e c o nd a p p l i c a t i o n. With b er r a i ns t h e re w as a g a in g r o w th of m a ny i n d i v i d u a l s. w i th s u ch p e r s i s t e n c e, we s e l e c t ed 350 c l o n es of good c o l o r, t u r e, e t c ., and g r ew t h em i n . f i e ld r o w s. L a te in t he s u m m er of 19 61, w h i le e v a l u a t i ng t h e s e, we n o t ed t h at e a ch o r i g i n al p l a nt f o r m ed a s m a ll t u ft f r om w h i ch s p r e a d i ng by s t o l o ns h ad t a k en p l a c e. F u r t h e r, we n o t ed t h at p l a n ts h ad s p r e ad into t h e ir own s h a de so t h at t h e re w as good s p r e a d i ng to t he n o r th and e a s t, but a l m o st no s p r e a d i ng to t he hot s o il of t he s o u th and w e s t. O c t o- I m p r e s s ed t e x- O c c a s i o n al e x c e p t i o ns w e re n o t ed a nd we f o u nd about 23 c l o n es t h at g r ew w e ll to t he s o u t h. We r e t e s t ed t h e se in r e p l i c a t ed a nd e n d ed with s ix t h at h ad a b i l i ty to s p r e ad into hot s o i l. As p u t t i ng g r e en t u r f, t h r ee w e re d i s c a r d ed f or d i s e a se a nd one f or an open, l o o se h a b it of g r o w t h. Of t he r e m a i n i ng we put o ne out to t e st w i th golf s u p e r i n t e n d e n t s. W i t h in two y e a r s, e v e ry t e st w as l o st d ue to c h a n g es in p e r s o n n e l, n ew c o n s t r u c t i on s u s c e p t i b i l i ty t w o, t r i a ls or s i m i l ar r e a s o n s. W h en we e x a m i n ed t he h i s t o ry we t h e re is a g r e at r e l u c t a n ce to u se g r a s s es t h at t i ve p l a n t i n g. r e a l i z ed r e q u i re v e g e t a- T h is is an e x a m p le of t he c o m m o n e st t y pe of g r a ss s e l e c t i on in t u rf i m p r o v e m e n t: p e r i or in one or m o re of i ts c h a r a c t e r i s t i c s. s i n g le p l a nt s e l e c t i on of an i n d i v i d u al s u- F a u l ts in o ur p r o g r am in 19 56 e v o k ed e n t h u s i a sm but i n c l u d ed d e v e l o p i ng a c l o ne w h en t h e re w as r e s i s t a n ce to v e g e t a t i ve p r o p a g a t i o n; l a ck of a d e- f i n ed p r o g r am w i th s t a t ed g o a l s; and s e l e c t i on t h at w as s t r o ng on q u a l i t i e s, w e ak on c o n d i t i o n s. T wo h u n d r ed n u r s e ry l o ts of C o n- g r e s s i o n a l, A r l i n g t on and C o h a n s ey b e n t g r a ss d i s t r i b u t ed to golf c o u r se s u p e r i n t e n d e n ts o n ly in v e g e t a t i ve p l a n t i ng of p a rt of one g r e e n. Q u a l i t i es vs c o n d i t i o ns is s h o wn in T a b le 1. An i m m a t u re p r o g r am m ay t e nd to n e g l e ct t he c o n d i t i o ns of t e s t, t h o u gh B u r t on h as c l e a r ly s t a t ed t h em in h is q u i d e l i n es ( B u r t o n, 19 69). I n f o r m al d i s c u s s i on at t he 1st I n t e r n a t i o n al T u r f g r a ss R e s e a r ch C o n f e r e n ce w as c o n c e r n ed w i th t he p o s s i b le d i s p a r i ty b e t w e en c o n d i t i o ns of t e st and c o n d i- t i o ns of u s e, a p r o b l em d i s c u s s ed by J u s ka a nd H a n s on ( 1962). Q u a l i t i es a re e m p h a s i z ed in r ow t e s t s, c o n d i t i o ns in plot t e s ts and t h e re is a t e n s i on b e t w e en t he two. A h l g r en et al. ( 1945) n o t ed t h at of 74 s u p e r i or c l o n es of 11,800 in f i e ld r o w s, only one r e m a i n ed s u p e r i or in a m o w ed plot. r e s u l t ed C o n d i t i on of T e s t i ng Q u a l t i t i es U n d er T e st T a b le 1 C l i m a te I r r i g a t i on M o w i ng F e r t i l i z a t i on S h a de C o m p e t i t i on C o l or ( e a r l y, m i d s e a s o n, l a t e) V i g or ( e s t a b l i s h m e n t, s p r e a d, r e c o v e r y) R e s i s t a n ce (to i n s e c t s, d i s e a s e s, n e m a t o d e s, w e e d s) T o l e r a n ce (to h e a t, cold, d r o u g h t, c l i p p i n g, w e a r) F o rm ( c o m p a ct of s p r e a d i n g) C o m p e t i t i v e n es s T e x t u re P e r s i s t e n ce R e g i o n a l i z ed G r a ss R e s e a r ch P r o g r a ms I c an f i nd only p r a i se f or t he m a ny t u rf i m p r o v e m e nt p r o- I s ay t h is b e c a u se r e g i o n al a d a p t a t i on of c o l o n i z i ng s p e- r o om g r a ms a c r o ss t he c o u n t r y, but I w o n d er if t h e re is not a l so f or s m a ll p r o g r a ms b a s ed on m a ss s e l e c t i on to m e et l o c al c o n d i- t i o n s. c i es is u s u al ( B r a d s h a w, 19 59). of s e v e r al E a s t e rn v a r i e t i es to p e r f o rm as w e ll in t he W e st as v a r i e t i es of W e s t e rn o r i g in and v i ce v e r s a; a l s o, m a ss d o es not r e q u i re t he e x p e n se of l a r ge r ow t r i a l s, r e s u l ts of w h i ch a re o f t en n e g a t ed in p l o t s. As an e x a m p le of r e g i o n al d i f f e r e n c e, n o te t he s u p e r i o r i ty of P e n n l a wn r ed f e s c ue in t he h u m id E a s t, t he i n d i f f e r e nt p e r f o r m a n ce of R a i n i e r; w h i le in t he hot d ry W e s t, R a i n i er is t he s u p e r i or g r a ss of t he two. I h a ve b e en i m p r e s s ed by f a i l u re s e l e c t i on We h a ve b e en s u f f i c i e n t ly i m p r e s s ed by t he s m a ll r e g i o n al t e st t h at we h a ve w r i t t en out a p r o g r a m, t h o u gh to d a te we h a ve not m o v ed to i m p l e m e nt it. S u ch a p r o g r am b a s ed on p o p u l a t i on g e n e t i c s, a l l o ws us to e m p h a s i ze t he c o n d i t i o ns of t e s t i n g, p a r- t i c u l a r ly t he m a ny c o n d i t i o ns of c l i m a te and c o m p e t i t i o n. T he m a n n er t h r o u gh d i s e a s e. F i r s t l y, m i ty of o ur g r e en c a r p et a nd m a k es p r e s e nt b l u e g r a s s es u n s u i t e d. S e c o n d l y, is in w h i ch we p r o p o se to get a h a n d le on t he p r o g r am it is d i s e a se t h at d e s t r o ys t he u n i f o r- is xiaAAixe oi V a r i g r a ss d i s e a s e s. T h e re a re t h r ee b a s ic p a r a s i t ic d i s e a s es of t u r f: r u s t, s m ut and m i l d e w. T he r e m a i n i ng d i s e a s es a re f a c u l a t i ve p a r a s i t es t h at a p p e ar only w h en t he g r a ss We p r o p o se t h en to u se as t he c o n d i t i o ns of t he t e s t, t h o se c o n- d i t i o ns w h i ch p r e d i s p o se to d i s e a s e, and to u se d i s e a se as t he s e l e c t i ve a g e nt to c a u se g e n e t ic s h i ft in o ur p o p u l a t i on ( w h e re d i s e a s es a re t h o se c a u s ed by t he f a c u l a t i ve p a r a s i t e s ). is s t r e s s ed to t he point of i n j u r y. T he s t r e ss f a c t o rs in o ur s i t u a t i on w h i ch l e ad to d i s e a se a re s t r e s s, c l i m a te ( h i gh t e m p e r a t u r e s ), s o il c o m p a c t i on and w a t er and t h o se f a c t o rs w h i ch i n c r e a se c o m p e t i t i on or r e d u ce c a r b o- h y d r a te s u p p l y, s u ch as f e r t i l i z a t i on and s h o rt a nd f r e q u e nt m o w- ing. A d v a n t a g es of R e g i o n al T e s t i ng P r o g r am T he a d v a n t a g es of s u ch a p r o g r am h a ve b e en n o t ed by H a n- In o r d i n a ry p o p u l a t i on g e n e t i cs one a s s u m es s on ( 19 72). B e g i n n i ng w i th a s e ed s o u r ce of h i gh g e n e t ic d i v e r s i t y, we q u i c k ly m a ke l a r ge g a i ns t o w a r ds o ur g o al w i th m i n i m um e f- f o rt and c o s t. t h at if a p r o p e r ty Z, h a v i ng a n o r m al d i s t r i b u t i o n, t h at t he m e an of t he p o p u l a t i on w i ll be c h a n g ed as in F i g u re 1, with t he m a x i m um s h i ft in t he f i r st s e l e c t ed g e n e r a t i o n. w o r k i ng with a p o p u l a t i on of a p o m i c t s, e s s e n t i a l ly a ll of t he s h i ft o c c u rs g e n e r a t i o ns would r e s u lt f r om i n c r e a s i ng t he s e l e c t i ve p r e s s u r e. V a r i a b i l i ty in s u b s e q u e nt g e n e r a t i o ns would r e s u lt as g e n e t ic n o i se a c c o m p a n y i ng s e l e c t i on p r e s s u re on o t h er c h a r a c t e r i s t i c s. If t he i n i t i al p o p u l a t i on is p a r t i a l ly a p o m i c t i c, s u b s e q u e nt g e n e r a- t i o ns m ay l o se in s e x u a l i ty and g a in in a p o m i x i s. in t he f i r st s e l e c t ed g e n e r a t i o n. B e n e f i ts in s u b s e q u e nt is s e l e c t ed f o r, If one is An a d d ed a d v a n t a ge of t he p r o g r am is t h at we would be s e- l e c t i ng f or r e s i s t a n ce to t he l o c al s t r a i ns of d i s e a se o r g a n i s m s. D i s a d v a n t a g es of R e g i o n al T e s t i ng P r o g r am D i s a d v a n t a g es of t he p r o g r am we h a ve o u t l i n ed a re s e v e r a l. is t h at it f a i ls to s e l e ct f or s u ch t u rf q u a l i t i es as c o l o r, F o r e m o st t e x t u re and d e n s i t y. T he l i t e r a t u re i n d i c a t es t h at b l u e g r a ss m ay be b r ed f or d i s e a se and i n s e ct r e s i s t a n c e, c l i m a t ic a d a p t a b i l i t y, t u rf h a b i t, s h a de t o l e r a n c e, t e x t u r e, d e n s i t y, c o l o r, r h i z o me e x t e n s i on and n u m b er e m e r g i n g. O ur p r o g r am s e l e c ts o n ly f or d i s e a se r e s i s t a n c e, t u rf h a b i t, c l i m a t ic a d a p t a b i l i ty and a g a i n st s e x u a l i ty ( H a n s on and J u s k a, 1962; Long, 1972). s e x u a l i t y, A s e c o nd d i s a d v a n t a ge is one to be c o n s i d e r ed in a ll p o p u l a- t i on w o r k - - r e p r o d u c t i ve p o t e n t i a l. T h e re is a l w a ys a d a n g er we w i ll l o se o ur g e n e t ic b a se by o v e r p r o d u c t i v i ty f r om a f ew i n d i- v i d u a l s, h e n ce s p e c i al t e c h n i q u es m u st be u s ed to s u p p r e ss e x- p r e s s i on of h i gh p r o d u c t i v i ty (we m i g ht h a r v e st e q u al s e ed w e i g h ts f r om e q u al a r e a s ). e l i m i n a t ed by s e l e c t i on m e an of t he n o r m al d i s t r i b u t i on m e an of p o p u l a t i on a f t er s e l e c t i on In t he f i r st s e l e c t ed g e n e r a t i o n, t he d i s- t r i b u t i on is a s s u m ed n o r m al about t he n ew m e a n. With a p o m i x i s, t he f i r st s e l e c t ed g e n e r a t i on h as a n on n o r m al d i s t r i b u- t i on about t he n ew m e a n. In u s i ng t he m e t h od of p o p u l a t i on g e n e t i cs to F i g u re 1. K e n t u c ky b l u e g r a s s, we a s s u me we a re s h i f t i ng t he p o p u l a t i on m e an of s o me c h a r a c t e r i s t i cs as i l l u s t r a t ed a b o v e. i m p r o ve A n o t h er d i s a d v a n t a ge is a l a ck of k n o w l e d ge of t he i n t e r a c- t i o ns b e t w e en t he f a c t o rs we a re s e l e c t i ng f o r, and of t he b a s ic a s- s u m p t i o ns we a re m a k i ng t h at m ay not be j u s t i f i e d; f or e x a m p l e, t h at c h a r a c t e rs do o c c ur in a n o r m al d i s t r i b u t i o n. A f i n al d i s a d v a n t a ge r e f e rs to o ur own r e q u i r e m e n t s. We h a ve l i m i t ed t i me and f u n ds and we r e q u i re t h at o ur w o rk y i e ld r e s u l ts t h at c an be g e n e r a l i z e d. Without s u ch r e s u l ts t h at p r o j e ct is one of s i m p le a p p l i c a t i on and s h o u ld be d o ne by i n d u s t ry r a t h er t h an t he u n i v e r s i t y. To o b t a in g e n e r a l i z a b le r e s u l ts r e q u i r es e x- t e n s i ve d a ta c o l l e c t i o n. O ur budget would j u s t i fy t h is only if we t h o u g ht t he r e s u l ts would be of v a l ue to oiir" c o n s t i t u e n c y. We d o n 't k n ow w h e t h er l o c al r e g i on is f e a s i b l e. W h i le w o rk h as b e en p l a n n e d, we p r o b a b ly s h a ll not e x e c u te it u n l e ss we f i nd a s t u d e nt who is p a r t i c u l a r ly e a g er f or s u ch a p r o g r a m, or u n l e ss f u r t h er d i s c u s s i on s h o ws how we c an a d v a n ce p o p u l a t i on s t u d i es by s u ch a p r o g r a m. l i m i t ed p r o d u c t i on of a v a r i e ty f or a In r e c e nt y e a rs I h a ve b e c o me i n t e r e s t ed not only in g r a s s e s, but in t he w h o le s u b c l a ss of m o n o c o ts ( M a d i s o n, 19 70). A m o ng p e r e n n i al c o l o n i z i ng s p e c i e s, w h i ch i n c l u d es m a ny m o n o c o ts (and d i c o t s ), a f o rm of a l t e r n a t i on of g e n e r a t i o ns e x i s t s. S e x u a l ly r e- p r o d u c i ng i n d i v i d u a ls a d a p t ed to a s i t u a t i on m ay r e p r o d u ce a s e x u a l ly o v er an i n d e f i n i te p e r i od of t i m e. s o me n u m b e rs m ay c h a n g e, and p o l y p l o i dy and a n e u p l o i dy m ay a p p e ar (Stucky and B a n f i e l d, 1946). S e x u al r e p r o d u c t i on is c o n- s e r v a t i ve and t e n ds to r e s t o re t he b a s ic g e n o m e. T o d ay I would f i nd t he m o st e x c i t i ng c h a l l e n ge in g r a ss g e n e t i cs t h at of i n v e s t i- g a t i ng t he g e n e t ic p l a s t i c i ty of a g r a ss as it r e p r o d u c es o v er d e- c a d es or c e n t u r i es by r h i z o m e s, t he a d a p t i ve v a l ue of t he c h a n g e s, and w h e t h er t h ey could be i n t r o d u c ed into b r e e d i ng p r o g r a m s. In t he a s e x u al p h a s e, c h r o m o- In an a p o m i ct s u ch as P oa p r a t e n s i s, we m ay a l r e a dy be t a k i ng a d v a n t a ge of g e n o me c h a n g e s. d e n ce s u g g e s ts t h at t he m u t a n ts a re l o st by s e x u al ( S t u c ky and B a n f i e l d, 1946). In A g r o s t is spp. t he e v i- r e p r o d u c t i on R e f e r e n c es C i t ed A h l g r e n, H. L., D. C. S m i t h, and E. L. N i e l s e n. 1945. B e h a v i or of v a r i o us s e l e c t i o ns of K e n t u c ky b l u e g r a ss ( P oa p r a t e n s i s) w h en g r o wn as s p a c ed p l a n ts and in m a ss s e e d l i n g s. J. A m. Soc. A g r o n. 37:268-281. B r a d s h a w, A. D. 19 59. P o p u l a t i on d i f f e r e n t i a t i on in A g r o s t is t e n u is Sibth., III. P o p u l a t i o ns in v a r i ed e n v i r o n m e n t s. N ew P h y t o l o g i st 59, 9 2 - 1 0 3. B u r t o n, G. W. 1969. I m p r o v i ng T u r f g r a s s e s. In T u r f g r a ss s c i e n c e. A m e r i c an S o c i e ty of A g r o n o my M o n o g r a ph #14. 410-424. H a n s o n, A. A. 19 72. B r e e d i ng of g r a s s e s. In T he b i o l o gy and u t i l i z a t i on of g r a s s e s. A c a d e m ic P r e s s, N. Y. pp. 3 7 - 5 0. H a n s o n, A. A., and F. V. J u s k a. 19 62. I n d u c ed m u t a t i o ns in K e n t u c ky b l u e g r a s s. C r op Sei. 2:369-371. J u s k a, F. V., and A. A. H a n s o n. 19 61. T he n i t r o g en v a r i a b le in t e s t i ng K e n t u c ky b l u e g r a ss f or t u r f. A g r o n. J. 5 3 : 4 0 9 - 4 1 0. Long, J. A. 1972. D e v e l o p i ng s u p e r i or t u rf v a r i e t i e s. In T he b i o l o gy and u t i l i z a t i on of g r a s s e s. A c a d e m ic P r e s s, N. Y. pp. 5 4 - 6 4. S ee a l so W e e d s, T r e es and T u rf 6(2): 12-14, 1967. M a d i s o n, J. H. 19 70. An a p p r e c i a t i on of m o n o c o t y l e d o n s. N o t es R o y. Bot. Gdn., E d i n b u r gh 30( 2 ) : 3 7 7 - 3 9 0. S t u c k e y, I. H., and W. C. B a n f i e l d. 1946. T he m o r p h o l o g i c al v a r i a t i o ns and t he o c c u r r e n ce of a n e u p l o i ds in s o me s p e c i es of A g r o s t is in R h o de I s l a n d. A m. J. Bot. 3 3 : 1 8 5 - 1 9 0. B R E E D I NG B E H A V I OR N O T ES ON S O ME I M P O R T A NT T U R F G R A SS S P E C I ES J o hn Long, E u g e ne M a y e r, V i r g il M e i e r, T e r ry R i o r d a n, Ming Yu B i o - C h e m i c al R e s e a r ch O. M. Scott & Sons C o m p a ny M a r y s v i l l e, Ohio 43040 I n t r o d u c t i on P r o g r e ss in i m p r o v i ng g r a s s es f or t u rf a p p l i c a t i o ns is i n f l u- e n c ed to a l a r ge d e g r ee by t he b r e e d i ng b e h a v i or of t he s u b j e ct g r a ss s p e c i e s. M e t h o ds e m p l o y ed f or c o m b i n i ng d e s i r ed t r a i ts in t he g r a s s es s e l e c t ed a re e f f e c t ed by a n u m b er of t he b r e e d i ng b e- h a v i or c h a r a c t e r i s t i c s. S o me of t he i m p o r t a nt b r e e d i ng b e h a v i or a s p e c ts i n c l u d e: (1) c h r o m o s o me n u m b er and p l o i dy l e v e l s; (2) m a g a s p o r o g e n e s i s; (3) m i c r o s p o r o g e n e s i s; (4) (5) self & c r o ss s t e r i l i t y / f e r t i l i t y; (6) a m p h i m i c t ic or a p o m i c t i c. e m b r y o g e n e s i s; P l a nt B r e e d i ng C h a r a c t e r i s t i cs C h r o m o s o me N u m b er and P l o i dy L e v e ls T he c h r o m o s o me n u m b e rs and p l o i dy l e v e ls of t he i m p o r t a nt t u r f g r a ss s p e c i es v a ry w i d e l y. W i t h in t he s p e c i es w h e re s e x u al r e p r o d u c t i on o c c u rs ( a m p h i m i c t i c ), c h r o m o s o me n u m b er w i ll h a ve a m a r k ed e f f e ct on t he m e t h o ds u s ed in b r e e d i ng and t he o u t c o m e. In g e n e r a l, h y b r i d i z a t i on of p a r e n t al t y p es w i t h in a s p e c i es with t he s a me c h r o m o s o me n u m b er is m o re l i k e ly to s u c c e ed t h an h y- b r i d i z a t i on of p a r e n t al t y p es with d i f f e r i ng c h r o m o s o me n u m b e r. L o ng and B a s h aw ( 1961) r e p o r t ed no s u c c e ss in c r o s s es m a de on s e l e c t ed St. A u g u s t i ne p a r e n t al t y p e s. A l a t er s t u dy of c h r o m o s o me n u m b e rs s h o w ed t h at o ne of t he p a r e n t al t y p es u s ed w as a t r i p l o i d, t h u s, a c c o u n t i ng f or l a ck of s u c c e ss i m b a l a n ce in t he t r i p l o id t y pe r e s u l t ed in a b n o r m al m i c r o s p o r o g e n e s is and m e g a s p o r o g e n e s i s. in h y b r i d i z a t i o n. C h r o m o s o m al T he e f f e ct of c h r o m o s o m al i m b a l a n ce h as b e en f o u nd to be m o st p r o n o u n c ed in i n t e r s p e c i f ic h y b r i d i z a t i o n. A l a ck of p r o p er p a i r i ng w h e re n u m b e rs v a ry and low or no h o m o l o gy with n e a r ly b a l a n c ed n u m b e rs l e ad to high s t e r i l i t y. In g r a ss s p e c i es w h e re a p o m i x is is p r e v a l e n t, v a r y i ng c h r o- m o s o me n u m b e rs h a ve not l e ad to h i gh s t e r i l i ty l e v e l s. M e g a s p o r o g e n e s is T h is r e p r e s e n ts t he s e q u e n ce of e v e n ts t h at l e ad up to t he r e p r o d u c t i on In g r a ss s p e c i es s u ch as L o l i um p e r e n n e, S t e n o t a p h r um d e v e l o p m e nt of t he egg c e ll in t he f e m a le p o r t i on of t he g r a ss f l o w e r. s e c u n d a t um and F e s t u ca r u b ra w h e re n o r m al s e x u al o c c u r s, a l i n e ar t e t r ad of s p o r es a re f o r m ed with one s p o re b e- c o m i ng f u n c t i o n al and t he r e m a i n i ng t h r ee d i s i n t e g r a t i n g. T he f u n c t i o n al m e g a s p o re t h en g i v es r i se to an eight n u c l e a te e m b r yo s ac as a r e s u lt of two s u c c e s s i ve m i t o t ic d i v i s i o n s. One n u c l e us of t he eight f u n c t i o ns as t he egg c e ll and f u s es with a n u c l e us f r om t he p o l l en t u be a f t er it e n t e rs t he e m b r yo s a c. A n o t h er n u c l e us f r om t he p o l l en t u be f u s es with p o l ar n u c l ei in t he e m b r yo s ac w h i ch t h en d e v e l o ps into e n d o s p e r m. T h is t h en r e p r e s e n ts t he s e q u e n ce of e v e n ts of m e g a s p o r o g e n e s is f or t he n o r m al s e x u al s p e c i e s. In a p o m i c t ic s p e c i e s, at l e a st t wo m e c h a n i s ms of a p o m i x is I n s t e ad of a l i n e ar t e t r ad of s p o r es f o r m i ng t he In c o n t r a s t, t he e m b r yo s ac of t he In P oa a l p i n a, P oa n e m o r a l i s, and P oa p a l u s t r is a re c o m m o n. d i p l o s p o ry o c c u r s. f r om an a r c h e g o n i al i n i t i al as t he c a se of s e x u al s p e c i e s, a r c h e g o n i al i n i t i al i t s e lf g o es t h r o u gh m i t o t ic d i v i s i o ns and f o r ms an u n r e d u c ed e m b r yo s a c. s e x u al s p e c i es h as a r e d u c ed c h r o m o s o me c o m p l e m e n t. A p o s- p o ry r e p r e s e n ts t he a p o m i c t ic m e c h a n i sm c o m m on to P oa p r a t e n- s is . e m b r yo s a c. n o r m al r e d u c ed egg m ay o c c u r. T h u s, v a r i ed c h r o m o s o me n u m- b e rs of s e x u al p r o g e ny m ay r e s u lt f r om f e r t i l i z a t i on of a r e d u c ed egg y i e l d i ng d i p l o i ds and t h o se p r o d u c ed f r om u n r e d u c ed eggs y i e l d i ng t r i p l o i d s. P r o g e ny m ay v a ry w i d e ly in c h r o m o s o me n u m b e r. In t h is i n s t a n ce a n u c e l l ar c e ll d e v e l o ps into an u n r e d u c ed In f a c u l t a t i ve a p o m i c ts s u ch as P oa p r a t e n s i s, a As one p l a ns a b r e e d i ng p r o g r a m, it is q u i te e s s e n t i al to t a ke i n to a c c o u nt t he e v e n ts in m e g a s p o r o g e n e s i s, p a r t i c u l a r ly w h e re a p o m i c t ic s p e c i es s u ch as P oa p r a t e n s is a re c o n s i d e r e d. M i c r o s p o r o g e n e s is W h e re m e g a s p o r o g e n e s is r e l a t es to t he f u n c t i o ns l e a d i ng to t he d e v e l o p m e nt of t he egg c e ll in what one would p e r h a ps c o n- s i d er t he f e m a le p h a s e, m i c r o s p o r o g e n e s is l e a d i ng to t he d e v e l o p m e nt of p o l l en w h i ch is c l a s s i f i ed as t he m a le p h a s e. T he s e q u e n ce s t a r ts f l o w e r. S p e c i al c e l ls ( m i c r o s p o re m o t h er c e l l s) u n d e r go r e d u c t i on and d i v i s i on to f o rm f o ur f u n c t i o n al m i c r o s p o r es w h i ch t h en m a t u re to f o rm p o l l en w i t h in t he a n t h e r s. g e n e s i s, r e d u c ed s p o r es a re f o r m e d. In t he s t e ps of m i c r o s p o r o- in t he a n t h e rs of t he g r a ss i n c l u d es t he e v e n ts C y t o l o g i c al s t u d i es of m i c r o s p o r o g e n e s is r e v e al s u b s t a n t i al i n f o r m a t i on to a l e rt t he p l a nt b r e e d er as to w h at m ay be e n c o u n- t e r ed in an i m p r o v e m e nt p r o g r am f or t he s p e c i es in q u e s t i o n. Long and B a s h aw ( 19 61) w e re a b le to d e t e r m i ne why h y b r i d i z a- t i on w as not s u c c e s s f ul in s e l e c t ed S t e n o t a p h r um t y p es a f t er c o m p l e t i ng a s u r v ey of m i c r o s p o r o g e n e s is of t he p a r- e n t al t y p e s. As r e p o r t ed a b o v e, one p a r e n t al t y pe w as A s t u dy of m i c r o s p o r o g e n e s is as would be e x p e c t e d. Without d e v e l o p i ng t h is i n f o r m a t i o n, c o n- s i d e r a b le t i me m ay h a ve b e en l o st in f u r t h er h y b r i d i z a t i on a t- t e m p t s. r e v e a l ed a h i gh d e g r ee of s t e r i l i ty s e c u n d a t um t r i p l o i d. E m b r yo g e ne s is F o l l o w i ng t he t wo p a r a l l el f u n c t i o ns of m e g a s p o r o g e n e s is and m i c r o s p o r o g e n e s i s, t he n e xt s e q u e n ce of e v e n ts l e a ds to t he d e- v e l o p m e nt of t he e m b r yo w h i ch r e p r e s e n ts t he n ew g e n e r a t i o n. T h is is a c r i t i c al p h a se in t e r ms of i n t r o d u c i ng d e s i r ed g e n e t ic v a r i a t i on into a p l a n n ed h y b r id c o m b i n a t i o n. p h a se is f o u nd in p o l l i n a t i on and f e r t i l i z a t i o n. L a ck of h o m o l o gy b e t w e en two p a r e n ts m ay be r e f l e c t ed in r e j e c t i on of t he d e s i r ed p o l l en by t he s t i g m a t ic s u r f a ce of t he e m b r yo s ac unit. c e p t a n ce o c c u r s, t he p o l l en g e r m i n a t es on t he s t i g m a t ic f a ce and f o r ms a p o l l en t u be w h i ch p e n e t r a t es into t he e m b r yo T he m o st s e n s i t i ve If a c- s u r- If c h r o m o s o m al i m b a l a n ce is a f a c t o r, t he f e r t i l i z a t i on s t ep s a c. m ay be b l o c k e d. D u r i ng p o l l en t u be g r o w th into t he e m b r yo s ac t wo m i t o t ic d i v i s i o ns o c c u r; w h e r e a s, d u r i ng s p e rm f o r m a t i on in t he a n i m al s y s t em no m i t o t ic d i v i s i o ns t a ke p l a c e. T h is s e n ts one of t he k ey d i f f e r e n c es in r e p r o d u c t i on in p l a n ts as c o n- t r a s t ed to r e p r o d u c t i on in a n i m a l s. P o l l en is m u ch m o re s i t i ve to c h r o m o s o m al i m b a l a n c es and d e f i c i e n c i es t h an s p e rm in a n i m a ls and as a r e s u lt t he p o l l en a b o r t s. T he egg c e l ls p l a n ts a re not as s e n s i t i ve as t he p o l l en to t he c h r o m o s o m al i m b a l a n c es and as a c o n s e q u e n ce t h e se i m b a l a n c es a re m o re l i k e ly to be t r a n s m i t t ed t h r o u gh t he f e m a le g a m e t e s. r e p r e- s e n- in A f t er t he p o l l en t u be p e n e t r a t es t he e m b r yo s a c, one n u c l e us f u s es w i th t he egg c e ll and a n o t h er f u s es w i th two p o l ar n u c l e i. T he f e r t i l i z ed egg c e ll t h en d e v e l o ps into t he e m b r yo w h i le t he o t h er f u s ed n u c l ei give r i se to e n d o s p e rm w h i ch f o r ms a p a rt of t he n ew s e e d. C h r o m o s o m al i m b a l a n c e, h o m o l o gy and g e n e t ic h o m o l o gy a re r e v e a l ed w h en s t u d i es a re m a de of t h e se e v e n t s. l a ck of c h r o m o s o m al In a p o m i c ts s u ch as P oa p r a t e n s i s, t he u n r e d u c ed e m b r yo s ac t he p o l ar n u c l ei to f o rm a nd r e s u l t i ng u n r e d u c ed egg d e v e l o ps into an e m b r yo without f u s i ng w i th t he n u c l e us a r i s i ng f r om t he p o l l en t u b e. T he n u c l e us f r om t he p o l l en t u be t h at f u s es w i th t he e n d o s p e rm h as b e en r e p o r t ed to s t i m u l a te f u r t h er d e v e l o p m e nt of t he u n r e d u c ed p r o - e m b r yo into an e m b r y o. In s p e c i es with s e x u al r e p r o d u c t i o n, s u ch as S t e n o t a p h r um s e c o n d a t um ( d i p l o i d s ), n o r- m al f u s i on of t he egg and t he n u c l e us f r om t he p o l l en t u be and t he o t h er t u be n u c l e us w i th t he p o l ar n u c l ei to f o rm e n d o s p e rm o c c u r s. T h is r e p r e s e n ts a s i g n i f i c a nt f u n c t i o n al d i f f e r e n ce b e t w e en a p o m i c ts a nd a m p h i m i c t s. Self & C r o ss S t e r i l i t y / F e r t i l i ty T he p l a n n i ng of a b r e e d i ng i m p r o v e m e nt p r o g r am m u st t a ke In t he s p e c i es C y n o d on d a c t y l o n, i n to a c c o u nt t h e se p h e n o m e n a. self s t e r i l i ty is q u i te c o m m o n. W h en c e r t a in br.eeding p r o c e d u r es a re u s ed to w o rk t o w a r ds plant u n i f o r m i t y, a h i gh d e g r ee of s t e r i l- i ty o c c u rs o b t a in a c c e p t a b le s e ed y i e l ds in A r i z o na C o m m on b e r m u d a g r a s s, In a v e ry p r a c t i c al s e n s e, l e a d i ng to low s e ed s e t. to « it is n e c e s s a ry to h a ve an u n r e l a t ed (to a d e g r e e) p o l l en s o u r ce to p r o v i de f or h i g h er c r o ss f e r t i l i z a t i o n. T h is l e a ds t h en to of u n i f o r m i ty in r e s u l t i ng p r o g e ny w h i ch is r e f l e c t ed in c o a r se t y p es n o t ed in l a wn p l a n t i n g s. l a ck Self s t e r i l i ty is a l so c o m m on in L o l i um p e r e n ne and t h at a p p r o p r i a te b r e e d i ng m e t h o ds be e m p l o y ed to c i r c u m v e nt self s t e r i l i ty p h e n o m e n a. r e q u i r es t he It is p a r t i c u l a r ly i m p o r t a nt to t a ke an i n v e n t o ry of t he a s p e ct of c r o ss c o m p a t i b i l i ty or i n c o m p a t i b i l i ty on n ew g r a ss p l a n n ed f or an i m p r o v e m e nt p r o g r a m. F or e x a m p l e, if one w e re to p l an an i m p r o v e m e nt p r o g r am on E r e m o c h l oa o p h u r o i d e s, a p r e r e q u i s i te would be to s t u dy t he c r o ss and self s t e r i l i ty or f e r- t i l i ty p h a se b e f o re p r o g r e ss m ay be r e a l i z e d. s p e c i es A m p h i m i c t ic or A p o m i c t ic P l a n ts t h at exhibit s e x u al r e p r o d u c t i on a re c l a s s i f i ed as a m- p h i m i c ts and t h o se t h at r e p r o d u ce by a s e x u al m e a ns a re c l a s s i- f i ed as a p o m i c ts ( G u s t a f s s o n, 1946). In t he a m p h i m i c t ic p r o c e s s, t he g a m e t o p h y te h as a r e d u c ed In t he g a m e t o p h y te an e g g - c e l l, t wo s y n e r- c h r o m o s o me n u m b e r. g i d s, t wo p o l ar n u c l ei and t h r ee a n t i p o d al c e l ls d e v e l o p. If t he e g g - c e ll w i th t he r e d u c ed c h r o m o s o me n u m b er in t h is p r o c e ss f o r ms an e m b r yo f o l l o w i ng f e r t i l i z a t i o n, t h is t h en s e x u al r e p r o d u c t i o n. P l a n ts in w h i ch r e p r o d u c t i on is not a s s o- c i a t ed w i th any f e r t i l i z a t i on a re t e r m ed a p o m i c t i c. r e p r e s e n ts In t he f i r st s u b g r o up ( a g a m o s p e r m y ), T wo s u b g r o u ps of a p o m i x is i n c l u de a g a m o s p e r my and v e g e- r e p r o- t a t i ve r e p r o d u c t i o n. r e- d u c t i on t a k es p l a ce by m e a ns of s e e d. r u n n e r s, p r o d u c t i on is by v e g e t a t i ve f o r m a t i o n s, s u ch as b u l b i l s, e t c. ( G u s t a f s s o n, 1946). T he t wo m e c h a n i s ms of a p o m i x is u n d er a g a m o s p e r my a re t e r m ed d i p l o s p o ry and a p o s p o r y. T h e se w e re d i s c u s s ed a b o ve u n d er m e g a s p o r o g e n e s i s. In t he s e c o nd s u b g r o u p, I m p o r t a nt t u rf s p e c i es in w h i ch a p o m i x is ( a g a m o s p e r m y) o c c u rs i n c l u de P oa p r a t e n s is and P a s p a l um n o t a t u m. T he a b o ve d i s c u s s i on on t he b r e e d i ng b e h a v i or a s p e ct w as not m a de in depth, but h o p e f u l ly c o v e r ed in s u f f i c i e nt d e t a il to e m- p h a s i ze t he i m p o r t a n ce of h a v i ng b a c k g r o u nd i n f o r m a t i on to p r o- v i de l e a ds on a p p r o p r i a te b r e e d i ng s y s t e ms to e m p l oy f or t u r f- g r a ss i m p r o v e m e n t. In t he f o l l o w i ng r e v i e w, s e v e r al i m p o r t a nt s p e c i es u s ed f or t u rf a p p l i c a t i o ns w i ll be c o n s i d e r ed r e l a t i ve to o ur k n o w l e d ge of b r e e d i ng b e h a v i or and p r o g r e ss t h at h as b e en m a de in a p p l i c a- t i on of a p p r o p r i a te b r e e d i ng s y s t e m s. T u rf S p e c i es B r e e d i ng B e h a v i or K e n t u c ky B l u e g r a ss ( P oa p r a t e n s i s) R e p r o d u c t i on in K e n t u c ky b l u e g r a ss is by p a r t i al ( f a c u l t a- t i v e) a p o m i x i s: T he a p o m i c t ic p r o c e ss a l l o ws r e t e n t i on of p a r t i c u l ar g e n o t y p es w h i ch u s u a l ly a re in a h e t e r o z y g o us s t a t e. In an i m p r o v e m e nt p r o g r a m, t he d e s i r ed g e n o t y pe is one e x h i b i- t i ng a h i gh d e g r ee of a p o m i x is f r om t he s t a n d p o i nt of v a r i e t al p u r i ty m a i n t e n a n ce in s e ed m u l t i p l i c a t i o n. T he i n f r e q u e n t ly o c c u r r i ng s e x u al r e p r o d u c t i ve p r o c e ss r . e l e a s es p o t e n t i al v a r i a b i l i t y, a l l o w i ng t he s p e c i es to a d a pt to d i f f e r e nt e n v i r o n m e n t al c o n d i t i o n s. T he p a r t i al s e x u al p r o- c e ss a l l o ws t he p l a nt b r e e d er to h y b r i d i ze p l a n ts w i th c o m- p l e m e n t a ry c h a r a c t e r i s t i c s. W h e re s e x u a l i ty is low and it is d e s i r ed to be i n c r e a s ed f or s e l e c t ed p a r e n ts f or b r e e d i ng p u r p o s e s, m u t a g e n ic a g e n ts s u ch as X - r ay h a ve b e en f o u nd e f f e c t i ve ( J u l e n, 1954). P o s s i b le m e t h o ds f or s p e c i es i m p r o v e m e nt i n c l u de s e l e c t i o n, h y b r i d i z a t i on and m u t a t i on b r e e d i n g. T he s e l e c t i on a nd h y b r i d i z a t i on m e t h o ds h a ve y i e l d ed i m p r o v ed In s e l e c t i o n, s u p e r i or p l a n ts a re t a k en f r om old t u rf a r e as v a r i e t i e s. or in s p a ce p l a n t i n g s. S u p e r i or p l a n ts a re e v a l u a t ed f or t u rf c h a r a c- t e r i s t i c s, f o l l o w ed by s e ed m u l t i p l i c a t i on and r e l e a s e. In h y b r i d i z a t i o n, h i g h ly a p o m i c t ic m a l es a re c r o s s ed to s u p- e r i or f e m a l es w i th a d e g r ee of s e x u a l i t y. S e l e c t i on m ay b e g in in t he Fi g e n e r a t i on b e c a u se m a t e r i al is h e t e r o z y g o u s, t h e r e f o r e, F^ p l a n ts l i k e ly a re d i f f e r e n t. S e g r e g a t i on in t he F2 g e n e r a t i on l i m i t ed b e c a u se of a p o m i x i s. is In m u t a t i on b r e e d i n g, s e l e c t i on m ay b e g in in M^ ( t he g e n e r a- t i on t h at w as t r e a t e d) b e c a u se of t he h e t e r o z y g o us c o n d i t i on of t he g e n o t y p e. S e g r e g a t i on is a g a in l i m i t ed in t he M 2 g e n e r a t i on b e c a u se of a p o m i x i s. C y t o g e n e t ic s t u d i es r e v e al c h r o m o s o me n u m b e rs r a n g i ng f r om 2n I 18 to 150 ( G r a z i, 19 61). C h r o m o s o me p a i r i ng m ay be d i s t u r b ed w h en p l a n ts a re s u b j e ct to v a r i ed c o n d i t i o ns ( G r u n, 1952). In s e ed f o r m a t i o n, c o m p e t i t i on b e t w e en a p o m i c t ic and s e x u al s e e ds d u r i ng e m b r yo f o r m a t i on m ay be i n f l u e n c ed by e n v i r o n m e n t. In c o m m e r c i al s e ed p r o d u c t i o n, l o c a t i on of f i e l ds in e n v i r o n m e n ts f a v o r a b le f or a p o m i c t ic s e ed f o r m a t i on s h o u ld be c o n s i d e r e d. In a i d i ng b r e e d i ng p r o g r a m s, m o re i n f o r m a t i on is n e e d ed on c o n d i t i o ns n e c e s s a ry f or s e ed f o r m a t i on s u ch as e n v i r o n m e n ts r e q u i r ed f or i n f l o r e s c e n ce i n i t i a t i on and f l o w e r i n g. S u ch i n f o r- m a t i on would aid h a n d l i ng p l a n ts with d i f f e r e nt m a t u r i t i es to c r o s s i ng w h i ch would a l l ow m o re f l e x i b i l i ty in t i me of y e ar w h en h y b r i d i z a t i on is c a r r i ed out. r e l a t i ve St. A u g u s t i n e g r a ss ( S t e n o t a p h r um s e c o n d a t u m) T h is s p e c i es is p r e s e n t ly u s ed e x t e n s i v e ly f or l a w ns in t he Gulf C o a st S t a t es and to a l i m i t ed e x t e nt in S o u t h e rn and C e n t r al C a l i f o r n i a. T he d i s t r i b u t i on of t h is s p e c i es f or l a w ns is l i m i t ed by t e m p e r a t u r e s. N o r m a l l y, it d o es not p e r s i st in r e g i o ns w h e re t e m p e r a t u r es r a n ge down to 10 F f or s e v e r al w e e k s. R e s e a r ch w as c a r r i ed out at T e x as A & M d u r i ng t he l a te 1950fs and e a r ly I 9 6 0fs to s t u dy b r e e d i ng b e h a v i or of St. A u g u s- t i n e g r a s s. F r om t h is r e s e a r ch ( L o ng and B a s h a w, 1961) it w as d e- t e r m i n ed t h at c h r o m o s o me n u m b e rs v a r i ed f r om 2n : 18 to 2n = 36. T he m e t h od of r e p r o d u c t i on w as f o u nd to be a m p h i m i c t i c. In d i p- l o id a c c e s s i o ns s t u d i e d, n o r m al m i c r o s p o r o g e n e s i s, m e g a s p o r o- g e n e s is a nd e m b r y o g e n e s is w e re o b s e r v e d. t i on s t u d i es r e v e a l ed t h at d i p l o id a c c e s s i o ns w e re c r o ss and self f e r t i l e. I n t r a s p e c i f ic h y b r i d i z a- P l a nt c o l l e c t i o ns o b t a i n ed f r om t he S o u t h e a s t e rn U n i t ed S t a t es w e re f o u nd to be l a r g e ly t r i p l o i d. C o l l e c t i o ns f r om t he W e s t e rn Gulf C o a st w e re f o u nd to be l a r g e ly d i p l o i d. C o l l e c t i o ns o b t a i n ed f r om A f r i ca w e re d i p l o id w h i le one a c c e s s i on o b t a i n ed f r om A u s- t r a l ia w as d e t e r m i n ed to be t r i p l o i d. I n t r a s p e c i f ic F^ h y b r i ds w e re f o u nd to be s u p e r i or in s o me c h a r a c t e r i s t i cs o v er p a r e n t al t y p es ( L o n g, 1972). T a ll F e s c ue ( F e s t u ca a r u n d i n a c e a) T a ll f e s c u e, a l t h o u gh c l a s s i f i ed as a c o o l - s e a s on p e r e n n i al r e- is c o n s i d e r ed one of t he m o st w i d e ly a d a p t ed g r a s s es It is u s ed f or t u rf r a n g i ng f r om t he S o u t h e rn g r a s s, l a t i ve to t e m p e r a t u r e s. t r a n s i t i on r e g i on to C a n a d a. It is a n a t u r a l ly c r o s s - p o l l i n a t ed g r a ss r a n g i ng f r om h i g h ly s e l f - s t e r i le to h i g h ly s e l f - f e r t i le ( C o w a n, 1956). A g r o n o m ic s t u- d i es in t he s p e c i es h a ve e m p h a s i z ed f o r a ge a p p l i c a t i o ns with a t t e n t i on g i v en to t u rf a p p l i c a t i o n s. T he a d a p t a b le b r e e d i ng m e t h o ds f or i m p r o v i ng t he s p e c i es a re c o n s i d e r ed as f o l l o w s: l i t t le 1. P h e n o t y p ic i n d i v i d u al s e l e c t i o n: B u r t on and D e v a ne ( 19 53) s h o w ed t h at high h e r i t a b i l i ty ( 7 6 - 9 0 %) as w e ll as wide g e n e t ic v a r i a b i l i ty (18-90%) e x i s ts f or c e r t a in c h a r a c t e r i s t i cs T h is i n d i c a t es t h at i n d i v i d u al s e l e c t i on to i m p r o ve t he s p e c i es f or t u rf a p p l i c a t i on is p r o b a b l e. S o u r c es of f r om a w i de a r ea of t he w o r ld w h e re it is g r o w n. I n d i v i d u al p l a n ts a re s e l e c t ed on t he b a s is of t h e ir own p e r f o r m a n ce f or one or m o re c h a r a c t e rs in s p a c e - p l a n t ed f i e ld n u r s e r i e s. T he s e l e c t ed p l a n ts a re u s ed e i t h er in c l o n al e v a l u a t i on or in c r o s s i ng w i th o t h er d e s i r- a b le m a t e r i al s h o u ld be t y p e s. in t a ll f e s c u e. i n c l u d ed 2. C o n t r o l l ed h y b r i d i z a t i o n: A l t h o u gh t he f l o r al o r g a ns of t a ll f e s c ue a re s m a l l, c e r t a in c o n t r o l l ed h y b r i d i z a t i on s t e ps c an be c a r r i ed out. By i n t e r c r o s s i ng s e l e c t ed p l a n ts and by p r a c- t i c i ng r e c u r r e nt s e l e c t i on w i t h in m a t e r n al l i n e s, an i n c r e a se t he f r e q u e n cy of d e s i r a b le g e n es m ay be w i ll p r o v i de n ew c o m b i n a t i o ns of c h a r a c t e r i s t i cs not s e en in t he p a r e nt and F^ p l a n t s. With d i a l l el c r o s s e s, b o th s p e c i f ic and g e n- e r al c o m b i n i ng a b i l i t i es of t he s e l e c t ed t y p e s, c an be e s t i m a t e d. in a c h i e v e d. T he F2 p o p u l a t i on 3. S y n t h e t ic v a r i e ty p r o c e d u r e: S y n t h e t ic v a r i e t i es a re m o re d e s i r a b le in t he b r e e d i ng of p o l y p l o id s p e c i e s, s u ch as t a ll f e s c u e. T he s e l e c t ed n o n - i n b r ed c l o n es c an be 2 to 6. T he p a r e n ts s e l e c t ed a re d e p e n d e nt m a i n ly on t h e ir high g e n e r al c o m b i n i ng a b i l i t i es and low i n b r e e d i ng d e p r e s s i o n. C l o n al p e r f o r m a n ce and s p e c i f ic c o m b i n i ng a b i l i ty r e p r e s e nt f u r t h er f a c t o rs f or c o n s i d e r a- tion. P o l y c r o s s es and s e l fs c an be u s ed to e v a l u a te g e n e r al c o m b i n i ng a b i l i ty and i n b r e e d i ng d e p r e s s i o n. C y t o g e n e t ic i n v e s t i g a t i o ns s h ow t he c h r o m o s o me n u m b er f or t a ll f e s c ue to be 2n z 42. M i c r o s p o r o g e n e s i s, m e g a s p o r o g e n e s is a nd e m b r y o g e n e s is a re t he s a me as f or o t h er s e x u a l ly r e p r o d u c ed p l a nt s p e c i e s. F i n e - L e a v ed F e s c u es ( F e s t u ca r u b r a, F e s t u ca ovina) T he f i n e - l e a v ed f e s c u es ( c r e e p i ng r ed f e s c ue and c h e w i n gs f e s c u e) a re u s ed w i d e ly in t he s a me g e n e r al r e g i on as K e n t u c ky b l u e g r a s s. V a r i e t i es of t h e se s p e c i es a re u s ed l a r g e ly in b l e n ds w i th K e n t u c ky b l u e g r a s s. M o st s e ed p r o d u c ts u s ed f or s h a d ed a r e as i n c l u de t he f i n e - l e a v ed f e s c u e s. T he b o t a n i c al v a r i e ty of F e s t u ca ovina v a r. d u r i u s c u la h as r e c e n t ly b e en i m p r o v ed by b r e e d i ng and f i n d i ng u se in t u rf s e ed p r o d u c t s. is C y t o g e n e t ic i n v e s t i g a t i o ns r e v e al c h r o m o s o me n u m b e rs of t he f i n e - l e a v ed f e s c u es f r om 2n z 28 to 2n z 56. T he b a s ic c h r o- m o s o me n u m b er of t he f i n e - l e a v ed f e s c u es is c o n s i d e r ed as s e v e n. T h e se s p e c i es a re c l a s s i f i ed as a m p h i m i c ts and exhibit n o r m al s e x u al r e p r o d u c t i o n. I m p r o v ed f i n e - l e a v ed f e s c u es h a ve b e en d e v e l o p ed by two b r e e d i ng m e t h o ds w h i ch a r e: 1. S e l e c t i o n: S e l e c t i on e i t h er m a ss or s i n g le l i ne t e s t ed f or t u rf q u a l i ty and s e ed y i e ld and t h en r e l e a s ed f or c o m m e r c i al a p p l i c a t i o n. 2. S y n t h e t ic v a r i e ty p r o c e d u r e: R e c e n t ly d e v e l o p ed v a r i e t i es a re t he r e s u lt of t h is b r e e d i ng p r o c e d u r e. L i n es or c l o n es by p o l y c r o ss t e c h n i q u es to m e a s u re c o m b i n i ng a b i l i ty a re s e l e c t ed f or u se in s y n t h e t ic v a r i e t i e s. t e s t ed T e s t i ng of i n d i v i d u al c o m p o n e nt c l o n es f or t u rf p e r f o r m a n ce a nd f l o w e r i ng a re c a r r i ed out in s y n t h e t ic v a r i e ty d e v e l o p m e nt p r o c e d u r e s. P e r e n n i al R y e g r a ss ( L o l i um p e r e n n e) R e n e w ed i n t e r e st in t he u se of p e r e n n i al r y e g r a ss f or f i ne t u rf a p p l i c a t i o ns no doubt h as r e s u l t ed f r om t he i n t r o d u c t i on of i m p r o v ed f i n e - l e a v ed t u rf t y pe v a r i e t i es w i t h in t he p a st f i ve y e a rs ( F u nk & E n g e l, 1968). I m p r o v e m e nt of p e r e n n i al r y e g r a ss h as r e s u l t ed t h r o u gh s y n t h e t ic v a r i e ty b r e e d i ng p r o c e d u r e s. O u t s t a n d i ng c l o n es a re t e s t ed by p o l y c r o ss t e c h n i q u es to a s s e ss c o m b i n i ng a b i l i t y. T h o se c o m b i n i ng w e ll a re s e l e c t ed f or i n c l u d i ng in t he m a k e up of s y n t h e t ic v a r i e t i e s. i m p r o v ed C y t o g e n e t ic i n v e s t i g a t i o ns s h ow p e r e n n i al r y e g r a ss to r e p r o d u ce s e x u a l l y. T he c h r o m o s o me n u m b er is 2n i 14 w i th a b a s ic c h r o m o- s o me n u m b er of s e v e n. S o me t e t r a p l o i ds h a ve b e en p r o d u c e d. B a h i a g r a ss ( P a s p a l um n o t a t u m) T h is s p e c i es is u s ed in t he S o u t h e a s t e rn U n i t ed S t a t es in lawn a p p l i c a t i o ns w h e re a u t i l i ty t y pe t u rf is d e s i r e d. t r a f f ic q u i te well, is f r ee of m a j or t u rf p e st p r o b l e m s, and is highly d r o u g ht It w i t h s t a n ds t o l e r a n t. I m p r o v ed v a r i e t i es h a ve b e en d e v e l o p ed by s e l e c t i ng o u t s t a n d- l a r g e ly ing c l o n es ( B u r t o n, 1951). f or f o r a ge a p p l i c a t i o n s. S u p e r i or c l o n e s, b a s ed on f o r a ge p e r f o r m a n c e, h a ve b e en i n c r e a s ed and r e l e a s e d. One v a r i e t y, A r g e n t i n e, a p p e a rs to be p r e f e r r ed f or t u rf a p p l i c a t i o n. I s o l a t ed c l o n es h a ve b e en t e s t ed Both a p o m i c t ic and s e x u al t y p es o c c ur w i t h in t he s p e c i e s. T he a p o m i c t ic t y p es a re c l a s s i f i ed as o b l i g a t e. R e f e r e n c es C i t ed B u r t o n, G. W. 1951. B r e e d i ng g r a s s es f or S o u t h e a s t e rn U n i t ed S t a t e s. Advan. A g r o n. 3, 197-240. B u r t o n, G. W., and E. H. D e V a n e. 19 53. E s t i m a t i ng h e r i t a b i l i ty in t a ll f e s c ue ( F e s t u ca a r u n d i n a c e a) f r om r e p l i c a t ed c l o n al m a t e r i a l. A g r o n. J o u r. 45:478-481. Cowan, J. R. 1956. T a ll f e s c u e. Advan. A g r o n. 3, 283-318. F u n k, C. R ., and R. E. E n g e l. 19 68. M a n h a t t an p e r e n n i al r y e g r a ss f or t u r f. T u r f g r a ss T i m es 4, No. 1. G r a z i, F ., M. U m a e r u s, and E. A k e r b e r g. 1961. O b s e r v a t i o ns on t he m o de of r e p r o d u c t i on and t he e m b r y o l o gy of P oa p r a t e n s i s. C o m m u n i c a t i o n s, S w e d i sh S e ed A s s o c. No. 133:489-541. G r u n, P. 19 52. E f f e ct of e n v i r o n m e nt u p on c h r o m o s o m al p a i r i ng of s o me s p e c i es and h y b r i ds of P o a. A m e r. J. Bot. 39 (5) 318-323. G u s t a f s s o n, A. 1946. A p o m i x is in h i g h er p l a n t s. P a rt 1. T he m e c h a n i s ms of a p o m i x i s. C. W. K. G l e e r u p, Lund. J u l e n, G. 1954. O b s e r v a t i o ns on X - r a y ed P oa p r a t e n s i s. A c ta A g r. Scand. 4, 585-59 3. Long, J. A., and E. C. B a s h a w. 1961. M i c r o s p o r o g e n e s is and c h r o m o s o me n u m b e rs 41-43. in St. A u g u s t i n e g r a s s. C r op Sci. 1, L o n g, J. A. 19 72. D e v e l o p i ng s u p e r i or t u rf v a r i e t i e s. In T he B i o l o gy and U t i l i z a t i on of G r a s s e s. A c a d e m ic P r e s s, N ew Y o r k. D i s c u s s i on P e r i od D r. W h a l l e y: H as t h e re b e en any w o rk done on t he i n d u c t i on of a p o m i x i s, in t r i p l o id S t e n o t a p h r um spp. so t h at t he p r o d u c t i on of v i a b le s e ed is p o s s i b l e? D r. L o n g: No. D r. D u i c h; H as a n y o ne e v er u s ed c o l c h i c i ne on St. A u g u s t i n e, p a r t i c u l a r ly in t h o se t r i p l o i d s? I w o r k ed s o me t i me with c o l c h i c i ne w h i le at T e x as A D r. Long: & M and f or s o me r e a s on we c o u l d n 't get it to w o r k. t a ke a n o t h er s p e c i es t h at w as s o m e w h at m o re d i f f i c u l t, t he d a l l a s- g r a ss ( P a s p a l um d i l a t a t u m ), c h r o m o s o m es up to 100, or d o u b le t h at one. be e v en m o re d i f f i c u lt to w o rk with. So f ar we h a v e n 't b e en s u c c e s s f ul w i th St. A u g u s t i n e. and w as a b le to t a ke it f r om 50 I did I t h o u g ht it would D r. M a d i s o n: Would you c h a r a c t e r i ze t he A f r i c an S t e n o t a p h r um s p p .? D r. Long: On t he A f r i c an b i o t y p e s, t h e re w e re two; one w as brought in f r om K e n ya and a n o t h er f r om South A f r i c a. T he one f r om K e n ya w as diploid. T he one f r om South A f r i ca w as an e x t r e m e ly c o a r se p l a nt and we w e re n e v er a b le to get f l o w e r i n g. t he r o ot t ip or s o m a t ic t i s s ue to c h e ck t h is out, but f r om t he g e n e r al a p p e a r a n c e, t e t r a p l o i d. T he one t e t r a p l o id t h at we p i c k ed up f r om t he F l o r i da c o l l e c t i on w as as you would e x p e ct r e l a t i ve to l a r g er p l a nt p a r t s. T he p o l l en w as s u b s t a n t i a l ly l a r g er t h an t he d i p l o i d s. it would h a ve a ll t he c h a r a c t e r i s t i cs of a I d i d n 't go to D r. D u d e c k: You c h a r a c t e r i z ed P a r a g u a y an and W i l m i n g t on as t e t r a p l o i d s, 40 c h r o m o s o me o b l i g a te a p o m i c t i c. Did you m e n- t i on A r g e n t i ne and P e n s a c o l a? D r. L o n g: is t he o b l i g a te a p o m i ct w i th 40 c h r o m o s o m e s. P e n s a c o la is t he diploid. A r g e n t i n e, as I u n d e r s t a n d, D r. P a y n e: in o r d er to s t i m u l a te s e ed d e v e l o p m e nt Is it n e c e s s a ry to h a ve p o l l en f r om an o u t s i de s o u r ce in an a p o m i c t? D r. F u n k: No, but a f o r e i gn p o l l en w i ll w o r k. D r. D a n i e l: W h a t 's t he c h r o m o s o me count of z o y s i a? D r. L o n g: Z o y s ia h as a c h r o m o s o me n u m b er of 2n - 40. P L A NT V A R I ET Y P R O T E C T I ON A CT AND T U R F G R A SS B R E E D I NG J o s e ph H i g g i ns USDA AMS G r a in D i v i s i on P l a nt V a r i e ty P r o t e c t i on O f f i ce 20782 H y a t t s v i l l e, M a r y l a nd I n t r o d u c t i on I would like to c o v er two i t e ms in t h is p r e s e n t a t i o n. One, d i s c u ss s o me of t he h i g h l i g h ts of t he P l a nt V a r i e ty P r o t e c t i on Act and, two, e x p l a in how we a re p r o c e s s i ng p l a nt v a r i e ty a p p l i- c a t i o n s. T he P l a nt V a r i e ty P r o t e c t i on Act w as s i g n ed into l aw on D e c e m b er 24, 19 70. l e g al p r o t e c t i on f or a p e r i od of 17 y e a rs to d e v e l o p e rs of n ew v a r i e t i es of p l a n ts w h i ch r e p r o d u ce by s e e d s. P r o t e c t i on h as b e en a v a i l a b le f or v e g e t a t i v e ly p r o p a g a t ed m a t e r i al t h r o u gh t he P a t e nt O f f i c e, U. S. D e p a r t m e nt of C o m m e r c e. C a r r o t s, c e l e r y, c u c u m b e r s, o k ra and p e p p e rs a re not i n c l u d ed in t he P l a nt V a r i e ty P r o t e c t i on Act. It e x t e n ds v o l u n t a r y, t o m a t o e s, R e q u i r e m e n ts f or P r o t e c t i on It m u st d i f f er f r om a ll k n o wn v a r i e t i es by T h r ee r e q u i r e m e n ts q u a l i fy n ew v a r i e t i es of p l a n ts f or p r o- t e c t i on as " n o v el v a r i e t i e s ." T he n o v el v a r i e ty m u st be d i s t i n c t, u n i f o rm and s t a b l e. at l e a st one i d e n t i f i a b le c h a r a c t e r. T h is could be m o r p h o l o g i c a l, p h y s i o l o g i c al or s o me o t h er c h a r a c t e r i s t i c, s u ch as t he baking q u a l i ty of a n ew v a r i e ty of w h e a t. Any v a r i a t i on t h at m ay e x i st in a v a r i e ty m u st be d e s c r i b a b l e, p r e d i c t a b le and c o m m e r c i a l ly a c- c e p t a b l e. F i n a l l y, t he v a r i e t y, w h en s e x u a l ly r e p r o d u c ed and r e c o n s t i- t u t e d, m u st r e m a in u n c h a n g ed in i ts e s s e n t i al and d i s t i n c t i ve c h a r- a c t e r i s t i cs to a d e g r ee e x p e c t ed of s i m i l a r ly d e v e l o p ed v a r i e t i e s. P r o t e c t i on c a n n ot be g i v en to a v a r i e ty if it h as b e en a p u b- in a t e c h n i c al j o u r n al m o re l ic v a r i e t y; i . e ., if it h as b e en f or s a le m o re t h an a y e ar in t h is c o u n t ry or if it w as d e s c r i b ed one y e ar b e f o re f i l i ng an a p p l i c a t i o n. O ur p r e s e nt l aw s t a t es t h at an a p p l i c a t i on w i ll not be a c c e p t ed a f t er one y e ar f r om t he f i l i ng d a te of an a p p l i c a t i on in a f o r e i gn c o u n t r y. H o w e v e r, we a re p r o p o s i ng to c h a n ge t he t i me p e r i od to not m o re t h an f o ur y e a r s. T he t i me p e r i od would be b a s ed on t e s t i ng r e q u i r e m e n ts of e a ch c o u n t r y. T he l aw is d e s i g n ed to p r o t e ct a n o v el v a r i e ty f r om b e i ng s o ld without t he c o n s e nt of t he o r i g i n a t or of t he v a r i e t y. t h an P r o c e d u r es N e c e s s a ry T he b r e e d er who w a n ts h is v a r i e ty p r o t e c t ed a p p l i es to t he t he i n d i c a t i ng S e c r e t a ry of A g r i c u l t u r e. T he b r e e d er m u st f u l ly d e s c r i be v a r i e ty b o t a n i c a l l y, i n c l u d i ng t he d i s t i n c t i ve f e s t u r es n o v e l t y, and give t he g e n e a l o gy and b r e e d i ng p r o c e d u r es oping it. An o b j e c t i ve d e s c r i p t i on f o rm to be f i l l ed out a l so w i ll f a c i l i t a te e s t a b l i s h m e nt of v a r i e t al n o v e l t y. A b r e e d er w i ll a l so be e x p e c t ed to d e p o s it and m a i n t a in v i a b le s e ed of h is v a r i e ty in a p u b l ic r e p o s i t o r y. T he a p p l i c a nt m u st a l so s t a te t he b a s is of h is o w n e r s h i p. in d e v e l- T he P l a nt V a r i e ty P r o t e c t i on O f f i ce w i ll m a i n t a in a of s c i e n t i f ic w o r ks to aid t he o f f i ce staff in a d m i n i s t e r i ng d u t i e s. A f t er c e r t i f i c a t es a re i s s u e d, s p e c i f i c a t i o ns of p r o t e c t ed p l a nt v a r i e t i es w i ll be p u b l i s h ed in t he O f f i c i al J o u r n al of t he P l a nt V a r i e ty P r o t e c t i on O f f i c e. F r om t i me to t i m e, as t h r o u gh an i n f o r- m a t i on p o r t i o ns of t he t e c h n o l o g i c al and o t h er p u b l ic i n f o r m a t i on a v a i l a b le to or w i t h in t he P l a nt V a r i e ty P r o t e c t i on O f f i ce to e n c o u r a ge n o v a t i on and p r o m o te t he p r o g r e ss of t he u s e f ul a r t s. s e r v i c e, t he O f f i ce w i ll d i s s e m i n a te to t he p u b l ic l i b r a ry t h e ir t h o se i n- We h o pe to g r a d u a l ly p r o v i de a v o l u n t a ry l i s t i ng u n d er o ur d a ta p r o c e s s i ng s y s t em f or t h o se v a r i e t i es d e v e l o p e d, but f or w h i ch no r i g h ts a re d e s i r e d. We w i ll t ry to h a ve as c o m p l e te a r e c o rd of a ll p l a nt v a r i e t i es T h is w i ll be done only as we c o m p l e te b a c k g r o u nd i n f o r m a t i on on e a ch s p e c i es f or s e a r ch p u r p o s e s, and w i ll a p p ly only to t h o se s p e c i es on w h i ch a p p l i c a t i o ns h a ve b e en f i l e d. in o ur o f f i ce f i l i ng s y s t em as is f e a s i b l e. O n ce t he v a r i e ty i n f o r m a t i on is c e n t r a l i z e d, m e a n i n g f ul and u s e f ul to t he o f f i ce and to a ll p l a nt s e a r ch and r e s e a r ch p u r p o s e s. it w i ll be u n i f o r m, b r e e d e rs f or T h e re a re f o r e i gn b r e e d e rs who would l i ke to r e c e i ve p r o t e c t i on u n d er o ur P l a nt V a r i e ty P r o t e c t i on Act and m a r k et t h e ir n o v el v a r i e t i es in t h is c o u n t r y. We p r o p o se to r e s t r i ct s u ch g r a n t i ng of c e r t i f i c a t es to t he e x t e nt t h at r i g h ts f or t he s a me s p e c i es a re g r a n t ed o ur n a t i o n a ls by t he c o u n t ry of t he a p p l i c a n t s. In o r d er to o r g a n i ze t he m a ss of d a ta d e s c r i b i ng m a ny c r op v a r i e t i es we a re d e v e l o p i ng a s e r i es of o b j e c t i ve d e s c r i p t i on f o r m s. T h e se f o r m s, w h en c o m p l e t e d, w i ll a l l ow us to f i le t he m a t i on in a c o m p u t e r i z ed d a ta p r o c e s s i ng s y s t em f or e a sy val. a c t e rs by w h i ch v a r i e t i es m ay be d i s t i n g u i s h e d. We h a ve e n d e a v o r ed to d e v i se m e a ns t h at w i ll b e st d e s c r i be or d e f i ne e a ch c h a r a c t e r. In d e v e l o p i ng t h e se f o r m s, we h a ve sought to i d e n t i fy c h a r- r e t r i e- i n f o r- T he b r e e d e rs a re u s u a l ly not c o n c e r n ed with a ll of t h e se c h a r- a c t e r i s t i cs and a re o f t en not e v en a w a re of s o me of t h e m. A n u m- b er of b r e e d e rs who h a ve v i e w ed s o me of t he f o r ms we h a ve d e v- e l o p ed in t he P l a nt V a r i e ty P r o t e c t i on O f f i ce h a ve s a id t h at b r e e d e rs would n ow r e c o g n i ze t he v a l ue in r e c o r d i ng m a ny of t h e se c h a r- a c t e r i s t i cs t h at t h ey h ad not r e c o r d ed r o u t i n e l y. And t h u s, f o r ms would h a ve t he e f f e ct of e n c o u r a g i ng b r e e d e rs m o re c o m p l e te d e s c r i p t i o ns of t h e ir n ew v a r i e t i e s. to m a ke t h e se W h en an a p p l i c a nt f or p l a nt v a r i e ty p r o t e c t i on f i l ls out t he o b j e c t i ve v a r i e ty d e s c r i p t i on f o r m, he is not r e q u i r ed to f i ll in a ll t he b l a n k s. H o w e v e r, we f e el it is in t he i n t e r e st of t he b r e e d er to f i ll in as m a ny b l a n ks as p o s s i b l e. t he F o r ms T he f o rm d e v e l o p ed f or b l u e g r a ss ( F i g. 1) i l l u s t r a t es a p p r o a ch we h a ve t a k en to d i s t i n g u i sh b e t w e en v a r i e t i e s. f or o t h er t u r f g r a ss g e n e ra w i ll be q u i te s i m i l ar to t h is one in m a ny r e s p e c t s. No a t t e m pt h as b e en m a de to m a ke a c o m p l e te d e s c r i p t i on f or e a ch v a r i e ty in t he o b j e c t i ve f o r m. H o w e v e r, t he f o rm s h o u ld a l l ow t he E x a m i n er to l i m it h is s e a r ch to a r e l a- t i v e ly f ew v a r i e t i es t h at m u st be c o m p a r ed f or o t h er e x p r e s s i o ns of d i f f e r e n c e. K e n t u c ky b l u e g r a ss c h a r a c t e r i s t i cs w e re r e l i ed on h e a v i ly in d e v e l o p i ng t he f o r m, s i n ce it is t he m o st b l u e g r a ss g r o wn f or t u r f. i m p o r t a nt T he p r i n c i p al c h a r a c t e r i s t i cs in t he f o rm a re g r o u p ed u n d er 18 h e a d i n g s. F or e x a m p le u n d er No. 6, Leaf B l a de -- c o l o r, s u r f a ce and s i ze m e a s u r e m e n ts a re i n c l u d e d. S t a n d a rd v a r i e t i es a re l i s t ed t h at r e p r e s e nt e a ch c o l o r. U n d er No. 9, P a n i c le -- s ix s t a n d a rd v a r i e t i es a re g i v en in o r d er to i n d i c a te t he n u m b er of c e n t i m e t e rs t h at t he n ew v a r i e ty is s h o r t er a n d / or P l a nt h e i g ht is i n d i c a t ed s i m i l a r ly w i th c o m p a r a t i ve v a r i e t i e s. l o n g er t h an one or two of t he s t a n d a rd v a r i e t i e s. T w e l ve i t e ms a re l i s t ed u n d er No. 17, E n v i r o n m e n t al A d a p- t a t i o n; f or e x a m p l e, h e a t, d r o u g ht and a ir p o l l u t i o n. An e x t ra b l a nk is p r o v i d ed to r e c o rd o t h er e n v i r o n m e n t al f a c t o r s. E a ch f a c t or l i s t ed is s c o r e d: 0 - Not t e s t e d, 1 - ' S u s c e p t i b l e, and 2 - R e s i s t a n t. U n d er No. 18, f o u r t e en of t he m o st i m p o r t a nt d i s e a s es of b l u e g r a ss a re l i s t ed t h at a re s c o r ed as g i v en u n d er No. 17. T he c h a r a c t e r i s t i cs i n c l u d ed in t he f o rm do not r e p r e s e nt t h at m ay e v e n t u a l ly be i n c l u d e d. With a d- a ll c h a r a c t e r i s t i cs d i t i o n al i n f o r m a t i on and e x p e r i e n ce in u s i ng t he o b j e c t i ve c r i p t i on f o r ms of d i f f e r e nt c r o p s, m o d i f i c a t i o ns w i ll be m a de t h e m. A d d i t i o n al c h a r a c t e r i s t i cs w i ll be i n c l u d ed and o t h e rs m o re a c c u r a t e ly or a p p r o p r i a t e ly d e f i n e d. A d d i t i o n al a n d / or o t h er c u l t i v ar s t a n d a r ds w i ll be d e s i r a b l e. C o l o rs m i g ht be s t a n d a r d i z ed with an a p p r o p r i a te c o l or s y s t em s u ch as N i c k e r s o n 's C o l or F a n. d e s- in t he We s o l i c it c o m m e n ts and c o n s t r u c t i ve s u g g e s t i o ns f or m o d i f y i ng t he f o rm to i m p r o ve on i ts p r a c t i c al u s a ge f or v a r i e- t al d e s c r i p t i o n. OBJECTIVE DESCRIPTION OF VARIETY Bluegrass (Poa spp.) Figure 1 1. KIND : l—i 1 = Poa compressa 3 = P. t r i v i a l is 2 = P. p r a t e n s is 4 = Other ( s p e c i f y) )—| Region of b e st a d a p t a t i o n: 1 = N o r t h e a st 3 = North C e n t r al 4 = P a c i f ic N.W. 2 = T r a n s i t i o n al Zone 5 = Other ( s p e c i f y) 2. MATURITY (At f i r st a n t h e s i s ): • Date: 1 = Early 3 = Medium (Newport) (Delta 1—i Number of days earlier than 1—; Number of days later than 2 = Medium early (Fylking) 4 = Late (Merion) . • . i—i 1 = Nugget 2 = Fylking 3 = Delta 4 = Merion 5 = Newport 6 = Baron 3. PLANT HEIGHT (Longest shoot 1 i i cm. Height 1 \ i cm. T a l l er than . I | j cm. S h o r t er than . . . Q . q 4. HABIT: 1 = Prostrate (Fylking) CZJ 3 = Erect (Delta) from s o il s u r f a ce to top or h e a d ): 1 = Nugget 3 = Delta 5 = Newport 2 = Fylking 4 = Merion 6 = Baron 2 = Semi-prostrate (Merion) 5. A VEGETATIVE REPRODUCTION Rhizomes Q Stolons (1 = Absent, 2 = Present): 6. LEAF BLADE : 1—i Color: 1 = Light Green (Rough Bluegrass) 2 = Moderately Dark Green (Merion) 3 = Dark Green (Adelphi) 4 = Blue Green (Canada Bluegrass) 5 = Other (specify) l—i Upper surface I—i Lower surface 1 = Shiny 2 = Dull 1 | | m m. Width i i \ m m. Length 7. LEAF SHEATH (Base): I—f Color ( S e e d l i n g ): S u r f a c e: q 1 = Glabrous 1 = Green 2 = Red 2 = Pubescent s i z e: i—i Keel: ' I—1 1 = Non -glaucous i Length 1 = Not Keeled | 2 = Keeled 2 = Glaucous 1 = Smooth 2 = Rough 8. LEAFINESS (At f i r st I—j Number of leaves per a n t h e s i s ): t i l l er or s h o o t: 1 = Few (1-3) (4-6) 2 = I n t e r m e d i a te 3 = Many (More than 6) 9. L PANICLE: 1—\ mm. Length 1—i mm. Longer than . • • • • 1—i mm. S h o r t er than . . . Q * = Nugget 3 = Delta 5 = Newport 2 = Fylking 4 = Merion 6 = Baron 1—| Number of p a n i c l es per P l a nt { | { Milligrams seed per p a n i c le Branches LOWEST WHORL: I—i 1 = Drooping (Prato) PANICLE HABIT: 1 = Nodding ( ) I—\ i—i mm. S p i k e l et Length • 2 = H o r i z o n t al (Merion) 3 = Other s p e c i fy 2 = Upright (Nugget) 10. LEMMA: i—| Basal Webbing: 1 = None 2 = Scant 3 = Copious Pubescence 1 = Glabrous 3 = Pubescent 2 = S l i g h t ly Pubescent 4 = Other q L a t e r al Nerves ( s p e c i f y) l—j Keel f ^j I n t e r m e d i a te Nerves l—i I n t e r m e d i a te Nerves: 1 = D i s t i n ct 2 = Obscure 11. SEED: i—i Apomictic 7oi I—| Phenol R e a c t i o n: S i z e: Weight: . [ ; \ i \ \ \ 1 • 1 ] Chromosome Number 1 = more than 95 2 = 85 to 95 3 = l e ss than 85 4 = Black (Delta - 2 hours) 2 = Beige (Cougar) (Merion) 1 = None - Lemma removed 3 = Brown (Windsor) 5 = Black (Anheuser - 24 hours) mm* Width [^j . j | • ; Grams per 10,000 seed (2n) mm. Length 12. (—[ 1 = Poor TURF DENSITY MAINTENANCE AT ONE INCH CUT): 2 = Moderate 4 = E x c e l l e nt 3 = Superior (Nugget) (Merion) 13. • VERTICAL GROWTH RATE: 1 = Slow (Nugget) 3 = F a st (Delta) 2 = Medium (Merion) 4 = Other ( s p e c i fy r e l a t i on to a standard) 14. SPRING GREEN UP: I—i 1 = E a r ly 2 = Medium 3 = L a te 15. FALL DORMANCY: • Northern • Southern (1 = Dormant (42°30! + 30f L a t .) (37°30f + 301 L a t .) 2 = I n t e r m e d i a te 1—| I n t e r m e d i a te 3 = Not Dormant) (40° + 301 L a t .) 16. SEEDLING VIGOR: 1 = Slow 1—| Seedling (growth 2 = Medium r a t e) 3 = F a st 17. ENVIRONMENTAL RESISTANCE TO (0 = Not Tested 2 = R e s i s t a n t ): 1 = Susceptible Cool Temperature i—i Cool Heat 1—I Heat "" -1 -- I—j Poor F e r t i l i ty 1—I S a l i n i ty I—i Air P o l l u t i on (Winter c o l o r) •• Drought ^ id S o il I—j S o il Compaction I—l Cold ( I n j u r y) 18. DISEASE, INSECTS, AND NEMATODE RESISTANCE TO: I—) Shade I—I A l k a l i n i ty \—jPoor Drainage ;—i Other ( s p e c i f y) (0 = Not Tested S u s c e p t i b le 1 = R e s i s t a n t ): 2 = l—i Helminthosporium vagans I—\ H. dictyoides j—j Erysiphe graminis I—{ Fusarium nivale f—I Typula iotana j—1 Puccinia graminia j—\ Pythium ultimatum 1—\ Other ( s p e c i f y) î—\ H. sorokinianum I—} Rhizoctonia solani I—\ Ustilago striiformis I—j F. roseum j—\ Scelerotinia homeocarpa I—\ P. striiformis I—j Crambus bonifatelluns INSTRUCTIONS COLOR: N i c k e r s o nfs or any recognized c o l or fan may be used to determine p l a nt c o l o rs of the d e s c r i b ed v a r i e t y. D i s c u s s i on P e r i od I t h i nk one of t he b e st w a ys to get s o me of y o ur D r. D u i c h: m a t i o n, s u ch as v a r i e t al t u rf p e r f o r m a n c e, would be t h r o u gh a c- c e p t ed r e g i o n al t e s t s. As f ar as t he o b j e c t i ve d e s c r i p t i on of a v a r i e t y, would it be a p p r o p r i a te f or y o ur o f f i ce to m a i n t a in a l o- c a t i on f or c e r t a in v a r i e t i es to r un r e f e r ee g r o w t h - t y pe t e s t s? i n f o r- D r. H i g g i n s: T h is is not n e c e s s a r y. T he b r e e d er m u st p r o v i de t he d a ta t h at w i ll c o n v i n ce t he E x a m i n er t h at h is v a r i e ty is d i s- t i n ct and m e e ts a ll t he r e q u i r e m e n t s. A lot of t h o u g ht h as b e en g i v en to m a k i ng c u l t i v ar d i s t i n c t i o ns w i t h o ut g r ow out t e s t s. We a re going to t ry t h is r o u te f i r s t. I t h i nk we c an t e ll b e t t er how o ur s y s t em w i ll w o rk a f t er we h a ve u s ed it a f ew y e a r s. D r. D a n i e l: T r oy is not in m a ny t e s ts or in m u ch p r o d u c t i o n. w o u ld s u g g e st s u b s t i t u t i ng N e w p o rt f or T r oy as one of t he s t a n d a rd v a r i e t i es b e c a u se of i t!s w i de u s e. I D r. H i g g i n s: Well, I am s u re you h a ve m a ny c o m m e n ts a nd a ll you n e ed to do is put t h em on t he f o rm d i s t r i b u t ed h e r e. P l e a se do not w a it u n t il 19 73 to m a ke y o ur s u g g e s t i o n s. Within t he n e xt f ew w e e ks w i ll be f i n e. a f t er you h a ve s e nt t h e se f o r ms in, p l e a se let me h e ar f r om you. If you get a n o t h er i d ea l a t e r, like t h is I b e l i e ve t h at " w h e re t he d a ta w as t a k e n" could be D r. D a n i e l: a d d ed to t he s e c t i on on s e ed h e ad h e i g h t. on w h e t h er t he s e ed w as g r o wn in I n d i a na or t he N o r t h w e s t, f or e x a m p l e. A l so w h e re you o f t en h a ve a c h o i ce b e t w e en n u m b e rs c o u ld e x p a nd t h is to at l e a st b r e e d er m u ch l e e w ay if he is only c h o o s i ng t he e x t r e m es or m i d d l e. -- one, two, t h r ee - - it s e e ms it would be good if you f i ve t h r o u g h o u t. You d o n 't give t he T h e re is a d i f f e r e n ce t h r ee t he D r. Long: D r. H i g g i n s, h a ve you s e en t he c h e ck l i st t h at D y v e n d ok u s e s? D r. H i g g i n s: Yes. w o r k? I w o n d e r ed if he h as f i n a l ly p u b l i s h ed t h is Dr Long: u s ed by t he IVRO in H o l l a n d. I t 's D y v e n d o k 's c h e c k l i st on m o r p h o l o g i c al f e a t u r es D r. H i g g i n s: t i o n ed s o me of t h e m. It is j u st l i m i t ed to c e r t a in o n e s, and I h a ve m e n- I t h i nk t h e re a re q u i te a f ew of t he c h a r a c t e r i s t i cs D r. Long: t h at m a t ch up with what D r. N i t t l er h as o b s e r v ed in s h e a th c o l o r. N ow he g o es into t he h a i rs and t he c o l or on t he s h e a t h. D r. D a d e, on o ur staff in O r e g o n, f i n ds he d o es not a g r ee with t he h a ir c h a r a c t e r i s t i cs t h at D y v e n d ok h as and h e 's c o me up w i th s o me r e f i n e m e n ts t h at we c o u ld p r o b a b ly p r o v i d e. D r. H i g g i n s: f r om you. T h is is t he v e ry t h i ng t h at I would l i ke to o b t a in D r. D u i c h: Would you o u t l i n e, by n a m e, t he t a b le of o r g a n i z a t i on of y o ur o f f i ce at t he p r e s e nt t i m e? I am s u re you k n ow t he two at t he t o p: Stan R o l l i n, D r. H i g g i n s: t he C o m m i s s i o n e r, a nd B e r n a rd L e e s e, t he Chief E x a m i n e r. My t i t le is E x a m i n e r. We h a ve f o ur o t h er E x a m i n e rs so f a r; H e r b e rt F i s h e r, K en E v a n s, Bob S n y d er and E l d on T a y l o r. T a y l or is a l so o ur L i b r a r i a n. S n y d er and F i s h er a re c o n c e r n ed with o r n a m e n t al p l a n ts and v e g e t a b l e s. I h a ve w o r k ed on p e as but I w i ll w o rk on a g r o n o m ic c r o p s, as w i ll K en E v a n s. T he L i b r a r i a n - E x a m i n er is o b t a i n i ng b o o ks and p e r i o d i c a ls f or o ur l i b r a ry and e n c o u r a g i ng p e o p le to s e nd in i n f o r m a t i on f r om a ll o v er t he c o u n t r y. We a l so h a ve two s e c r e t a r i e s. M r. M a y e r: What a re t he c o s ts i n v o l v ed w h en a p p l y i ng f or a p l a nt p a t e n t? T he a p p l i c a t i on f ee is $50. H o w e v e r, as s o on as D r. H i g g i n s: t he r e g u l a t i o ns a re a p p r o v e d, it w i ll be $250. T h e re w i ll be an a d d i t i o n al $250 c h a r ge to m a ke t he s e a r ch and $250 w h en a c e r- t i f i c a te is a w a r d e d. F i r s t, on t h is m a t u r i t y, I t h i nk b l o o m i ng m i g ht be D r. N i t t l e r: m u ch m o re p r e c i se t h an a h e a d i ng d a t e. S e c o n d l y, t h is q u i r e m e nt f or u n i f o r m i t y, t h at w i ll h a ve to be i n t e r p r e t ed p r e t ty b r o a d ly with t h is c r o ss p o l l i n a t ed c r o p, w o n 't i t? r e- D r. H i g g i n s: Yes. H o w e v e r, t h is w i ll d e p e nd on t he c r o p. D r. P a y n e: Would you b r i e f ly d e s c r i be t h is " s e a r c h" you s p e ak of a f t er you r e c e i ve t h is a p p l i c a t i o n. What is y o ur p r o c e d u re f r om t he t i me you r e c e i ve t h i s? I c a n n ot t e ll you in c o m p l e te d e t a il b e c a u se we D r. H i g g i n s: a re in t he p r o c e ss of d e v e l o p i ng a s e a r ch p r o c e d u r e, and I t h i nk it w i ll v a ry f or d i f f e r e nt c r o ps -- t he i n f o r m a t i on a v a i l- a b l e, and so on. We o b t a in d e s c r i p t i o ns of t he v a r i e t i es t h at a re in c o m m e r c e. We h a ve l o c a t ed a n u m b er of r e f e r e n c es t h at a re good. T h ey d e s c r i be s o me of t he o l d er v a r i e t i e s. F or m a n y, it w i ll be h a rd to get m u ch i n f o r m a t i o n. D r. P a y n e: Will t h is u l t i m a t e ly be c o m p u t e r i z ed in t he m a n n er of t he f i n g e r p r i nt s y s t em w h e re a b u t t on is p u s h ed and a s e l e c t ed f ew a re d e l i v e r ed f or s t u d y? T he i n f o r m a t i on s t o r ed in t he c o m p u t er w i ll c o me D r. H i g g i n s: f r om t he o b j e c t i ve d e s c r i p t i on f o r ms and, as I h a ve t h is i n f o r m a t i on w i ll not d e s c r i be a v a r i e ty c o m p l e t e l y. going to be a m e t h od to h e lp us z e ro in on a f ew r e l a t ed v a r i e t i e s. i n d i c a t e d, It is M r. S i m m o n s: h a n d l i ng m a t e r i al f r om f o r e i gn c o u n t r i e s? Could you r e v i ew y o ur p o t e n t i al p l a ns f or D r. H i g g i n s: We w i ll not a c c e pt an a p p l i c a t i on f r om a f o r e i gn c o u n t ry u n l e ss t h at f o r e i gn c o u n t ry r e c i p r o c a t es w i th s i m i l ar p r o t e c t i on to i n d i v i d u a ls f r om t he United S t a t es f or s p e c i f ic c r o p. is p r o t e c t ed in G e r m a n y, t h en we w i ll give a p p l i c a n ts f r om G e r m a ny p r o t e c t i on f or K e n- t u c ky b l u e g r a ss If K e n t u c ky b l u e g r a ss in t he U n i t ed S t a t e s. t h at D r. D u i c h: At t he p r e s e nt t i me w i th a g r o n o m ic c r o ps we h a ve r e g i s t r a t i on w i th T he A m e r i c an S o c i e ty of A g r o n o m y, as p u b- l i s h ed in C r op S c i e n c e. H a ve you p e o p le g i v en a ny c o n s i d e r a t i on to a ny t y pe of c o o r d i n a t i on of t h is r e g i s t r a t i o n? F or e x a m p l e, we get i n v o l v ed in a lot of d o u b le p a p er w o rk b e c a u se now, e x a m p l e, t h ey want p e r f o r m a n ce d a t a, a nd t h ey h a ve a c o m m i t t ee in m u l t i p le l o c a t i o n s. f or D r. H i g g i n s: We a re w o r k i ng c l o s e ly w i th C r op S c i e n c e, and t h is w h o le p r o g r a m, to c o o r d i n a te o ur w o rk with t h is g r o u p. t h at p l a nt v a r i e ty p r o t e c t i on h ad b e en I did t h is b a s ed on s o me of t he e x p e r i e n c es D r. F u nk D r. D u i c h: W h en I s u b m i t t ed P e n n f i ne f or r e g i s t r a t i on in C r op S c i e n ce I c o n t a c t ed Stan R o l l in and a s k ed if we could put a s t a t e m e nt on t he l a st l i n e, a p p l i ed f o r. h ad w i th M a n h a t t an p e r e n n i al r y e g r a s s? M a y be s c a re off s o me of t he s c a v e n g e r s. w as m o re or l e ss t he p r e c e d e nt f or he went a l o ng with it in t h is c a s e. Is t h is going to be a l l o w e d? I g u e ss t h is D r. H i g g i n s: A f t er you a p p ly f or a p r o t e c t i o n, you c an s t a rt s e l l i ng t he s e e d, T he s e ed l a b el m u st i n d i c a te t h at p r o t e c t i on h as b e en a p p l i ed f o r, and t h at w i t h o ut a r r a n g e m e nt w i th t he o w n e r. s e x u al r e p r o d u c t i on is not a l l o w ed D r. D u i c h: J u st like " p a t e nt a p p l i ed f o r ?" D r. H i g g i n s: Y e s. D r. D a n i e l: At what point w i ll you t a ke o v er f r om t he P l a nt P a t e nt O f f i c e. We h a ve p r e v i o u s ly w o r k ed with b l u e g r a s s es t h r o u gh t he P l a nt P a t e nt O f f i c e? D r. H i g g i n s: We a re a c c e p t i ng a p p l i c a t i o ns to p r o t e ct v a r i e t i es of K e n t u c ky b l u e g r a ss now. B a s ed on t he t i on of t he A c t, K e n t u c ky b l u e g r a ss w o r d i ng and i n c l u d e d. i n t e r p r e t a- is D r. D a n i e l: a g r e e m e n t. T h is h as not b e en s p e c i f i ed by law, or e v en by an I c an r e p ly to t h at a l i t t le bit. W h en t h ey w e re w o r k i ng D r. D u i c h; on d e v e l o p i ng t h is A c t, t h e re w as n o t h i ng s p e c i f i c a l ly s a id about b l u e g r a ss and of c o u r se e v en t he b l u e g r a ss p a t e nt b e i ng a w a r d ed u n d er t he P l a nt P a t e nt P r o t e c t i on Act is a l i t t le f u z zy or in t he g r ey a r e a, and I v e ry s p e c i f i c a l ly a s k ed A S TA to get b l u e g r a ss u n d er t h is r a t h er t h an t he 19 37 P l a nt P a t e nt A c t. T h ey s a id t he r e a s on t h ey did not want in t h e re s p e c i f i c a l l y, t h at s i n ce t h e re w as t h is m a t t er of v e g e t a t i ve r e p r o d u c t i o n, w as t h e re w e re o t h er c r o p s, p a r t i c u l a r ly in t he o r n a m e n t al a r e a. If t h ey t h ey c o v e r ed a n y t h i ng p e r t a i n i ng to a s e x u al r e p r o d u c t i o n, w e re going to get a lot of s t a t ic on t he Act i t s e l f. In o r d er to in e l i m i n a te a ll t h is o p p o s i t i o n, t h ey d r o p p ed b l u e g r a ss p er s e, as f ar as n a m i ng it in t he P l a nt V a r i e ty P r o t e c t i on Act of 19 70. to put b l u e g r a ss D r. D a n i e l: And y e t, D r. H i g g i n s, you h a ve s o me a p p l i c a t i o ns f or b l u e g r a s s es a l r e a d y? D r. H i g g i n s: Y e s, s e v e n. H o w e v e r, I d o n 't k n ow if t h e se a re v e g e t a t i v e ly or s e x u a l ly r e p r o d u c ed as to w h e t h er t h ey w i ll get p r o t e c t i on or not. In d i s c u s s i o ns b a ck in J a n u a ry w i th Stan R o l l i n, I D r. L o n g: t h i nk a ny b l u e g r a ss t h at is p r o p a g a t ed by s e ed w i ll fit t he P l a nt V a r i e ty P r o t e c t i on Act. A c a se l i ke W a r r e n 's v e g e t a t i v e ly p r o p a g a t ed b l u e g r a ss would h a ve to go u n d er t he P l a nt P a t e nt A c t. We would p l an to h a n d le t he St. A u g u s t i ne and b e r m u d a- g r a s s es t h at a re p r o p a g a t ed s t r i c t ly v e g a t i v e ly u n d er t he P l a nt P a t e nt A c t, and not t he P l a nt V a r i e ty P r o t e c t i on Act. So t he l a n g u a ge in t he 19 70 a ct is s u f f i c i e n t ly b r o ad to t a ke t he b l u e- g r a s s e s, p r o v i d ed you p r o p a g a te t h em by s e e d. D r. P a y n e: You m e n t i o n ed p a t e n t i ng by a f o r e i gn c o u n t r y. What a b o ut O. M. Scott & S o n s, f or e x a m p l e, a p p l y i ng f or t he p a t e nt of a l i ne f r om a f o r e i gn c o u n t r y? M r. D o e l l i n g e r: We j u st a t t e n d ed a f o r um s p o n s o r ed by t he A m e r i c an S e ed T r a de A s s o c i a t i on l a st w e ek m P o r t l a nd and it w as s t a t ed t h at a U. S. N a t i o n al m ay o b t a in p r o t e c t i on of a n o v el v a r i e ty e v en t h o u gh it r e s u l ts f r om b r e e d i ng and r e s e a r ch c o n d u c t ed in a f o r e i gn c o u n t r y. We u n d e r s t a nd f u r t h e r, t h at a N a t i o n al of t he f o r e i gn c o u n t ry m ay a s s i gn t he o w n e r s h ip of a n o v el v a r i e ty to a N a t i o n al of t he U n i t ed S t a t es who m ay go a h e ad and a p p ly f or p r o t e c t i o n. D r. Yu: D r. H i g g i n s, we k n ow t h at b l u e g r a ss v a r i a t i on f or c h r o m o s o me n u m b e rs e x i st a m o ng v a r i e t i e s. c a se a p a t e n t ed v a r i e ty h ad p r o d u c ed s o me p r o g e n i es w i th a d i f- f e r e nt c h r o m o s o me n u m b e r, but w as not d e t e c t ed by t he o r i g i n al p a t e n t e e, can a n o t h er p e r s on who d e t e c t ed t h is c h a n ge in c h r o- m o s o me n u m b er a p p ly f or a n ew p a t e nt on t h is p l a n t, a l t h o u gh t he o t h er p l a nt c h a r a c t e r i s t i cs r e m a in t he s a m e? is a p o m i c t ic and In D r. H i g g i n s: A c c o r d i ng to t he Act, t he p l a nt cannot c h a n g e. T h at is one of t he c h a r a c t e r i s t i cs of u n i f o r m i t y. d i s t i n g u i s h i ng c h a r a c t e r, t he o w n er would not h a ve p r o t e c t i o n. T he o w n er would h a ve to s u b m it a n ew a p p l i c a t i o n. If t h is w e re t he only D r. Long: One s u g g e s t i on t h at I t h i nk we s h o u ld c o n s i d er on t he f o rm f or b l u e g r a s s es would be to u se a p p r o x i m a te f or c h r o m o s o me n u m b e r. D r. H i g g i n s: t h at r a n g e. P o s s i b ly so, but t he v a r i a b i l i ty would h a ve to be in It could not v a ry any m o re t h an t h at r a n g e. To i l l u s t r a te t he a p p r o x i m a t e, t he S w e d es j u st count in b l u e- g r o up it to t he n e a r e st D r. D a n i e l: to t he n e a r e st n u m b er of s e v e n. T h ey s ay c h r o m o s o m es g r a ss t e nd to r un in s e v e ns so t h ey j u st m u l t i p le of s e v en as a m e a ns of e x p e d i t i ng t h e ir o b s e r v a t i o n al w o r k. D r. H i g g i n s: T h is h as a p p l i c a t i on to o t h er c r o p s. M a y be w ay to h a n d le it i s, i n s t e ad of a s k i ng f or t he 2n n u m b er of c h r o m o s o m es b e r. T h at would c o v er t he s a me t h i n g, would it n o t? t h at would only be one f a c t o r. in t he f o r m, t h ey s h o u ld give at l e a st t he n n u m- t he I t h i nk D r. D a n i e l: How s o on do you h o pe to f i n a l i ze t h is f o r m? You h a ve s t a r t e d; you h a ve a c e r t a in b e g i n n i ng h e r e. D r. H i g g i n s: g e s t i o ns f or i m p r o v i ng t he f o r m. I w i ll w o rk on it as s o on as I r e c e i ve y o ur s u g- I would s ay w i t h in t wo m o n t h s. I a s s u me we s u b m it d a ta as an a d d e n d um to t h is M r. M a y e r: f o r m. Will t h e re be a f o r m at to u se on t he t y pe of d a ta w e ' re s y p p o se to s u b m i t? t h at D r. H i g g i n s: T h e re a re no s p e c i al d e t a i ls in t h at r e g a r d. B R E E D I NG S E E D ED V A R I E T I ES OF B E R M U D A G R A SS F OR T U R F G R A SS USE W i l l i am R. K n e e b o ne D e p a r t m e nt of A g r o n o my and P l a nt G e n e t i cs U n i v e r s i ty of A r i z o na 85721 T u c s o n, A r i z o na I n t r o d u c t i on T he bulk of t he b e r m u d a g r a ss s e ed m o v i ng in w o r ld t r a de f or t he p a st half c e n t u ry h as b e en p r o d u c ed in A r i z o n a. As a l- f a l fa s e ed p r o d u c t i on in t he Y u ma a r ea d e c l i n ed u n d er a c o m b i n- a t i on of f e w er p o l l i n a t o rs and i n c r e a s ed s o il s a l i n i t y, b e r m u d a- g r a s s, p r e s e nt as a w e e d, i n c r e a s ed u n d er s a l i n i ty l e v e ls h i gh f or t he a l f a l f a, and b e r m u d a g r a ss s e ed c h a n g ed f r om a b y- p r o d u ct to a c a sh c r o p. T he d r a w b a c ks and s o me of t he a d v a n- t a g es of A r i z o na b e r m u d a g r a ss s e ed f or t u rf r e l a te to c y c l ic s e ed r e p r o d u c t i on u n d er i r r i g a t i o n; f i r st as a w e ed and l a t er as a c r op in t he hot, d ry c l i m a te at Y u m a. T he f i r st b e r m u d a g r a ss in t he a r ea p r o b a b ly a r r i v ed a c e n t u ry or so a go f r om W e s t e rn M e x i co and f r om S o u t h e rn C a l i f o r n ia ( K n e e b o n e, 1966). C h a r a c t e r i s t i cs r e l a t ed to o r i g i ns and h i s t o ry i n c l u d e: s a l ty c o n d i t i o ns and m i s m a n a g e m e n t, s u s c e p t i b i l i ty to d i s e a s es u n d er h u m id c o n d i t i o n s, r a p id s p r e ad and l a ck of w i n t e r h a r d i n e s s. t oo c a p a c i ty to e n d u re d r o u t h, A f r e q u e nt c o n t a m i n a nt in b e r m u d a g r a ss s e ed is giant b e r- m u d a g r a s s, c o m m on in t he Y u ma a r ea and p r o b a b ly s t e m m i ng f r om i n t r o d u c t i o ns f r om I n d i a, t e s t ed t h e re by t he B u r e au of R e- c l a m a t i on b e f o re W o r ld W ar I. N a t u r al c r o s s i ng f r e q u e n t ly o c- c u rs b e t w e en t he d i p l o id giant b e r m u d a g r a ss and t he t e t r a p l o id b e r m u d a g r a s s. Giant b e r m u d a g r a s s, as i ts n a me i m p l i e s, c o a r s e, u n d e s i r a b le as a t u rf and it is m u ch l e ss t o l e r a nt of S o u t h e a s t e rn p a t h o g e ns t h an b e r m u d a g r a s s. is B e r m u d a g r a ss R e s e a r ch P r o g r am T he p r e s e nt b e r m u d a g r a ss r e s e a r ch p r o g r am in A r i z o na b e- g an in 19 59 in r e s p o n se to c o n c e rn by s e ed g r o w e rs with p o t e n t i al m a r k et l o s s es to v e g e t a t i v e ly p r o p a g a t ed v a r i e t i e s. A c e r t i f i c a- t i on p r o g r am w as d e v e l o p ed to p r o v i de S o u t h e rn c o n s u m e rs a s- s u r a n ce t h at t h ey would h a ve no c o n t a m i n a t i on f r om giant b e r m u- d a g r a ss and c o l l e c t i o ns of b r e e d i ng m a t e r i al w e re a s s e m b l ed to a s s e ss t he p o t e n t i a ls and b e g in an i m p r o v e m e nt p r o g r a m. A r i- z o na c e r t i f i ed s e ed t o d ay is not only f r ee of giant b e r m u d a g r a s s, but is c o n s i d e r a b ly m o re u n i f o rm in t y p e, a r e s u lt of s t r i n g e nt r o g u e i ng of o f f - t y p es in t wo c y c l es of r e p r o d u c t i on of t y p i c al b e r- m u d a g r a ss p r e c e d i ng p r e s e nt f o u n d a t i on s e ed p r o d u c t i o n. T h is s e ed m u st be u s ed to e s t a b l i sh n ew c e r t i f i ed s e ed f i e l d s. As yet, h o w e v e r, a d i s t i n c t ly d i f f e r e nt s e e d ed t u rf t y pe v a r i e ty h as not b e en r e l e a s e d. i n- H a r l an et al. (19 70) d e s c r i be eight s p e c i es and t en b o t a n i c al v a r i e t i es of C y n o d on f r om w h i ch t h r ee of t h e ir s ix v a r i e t i es of C. d a c t y l on ( d a c t y l o n, e l e g a n s, p o l e v a n s i i ), both v a r i e t i es of C. c o m p l e t us ( i n c o m p l e t us and h i r s u t u s) and C. t r a n s v a a l e n s is c an be c o n s i d e r ed to h a ve t u r f g r a ss p o t e n t i a l. T he v a r i e t i es of C. d a c t y- l on a re t e t r a p l o id (2n - 36), t he o t h e rs diploid (2n - 18) with o c- c a s i o n al t e t r a p l o i ds in C. i n c o m p l e t us v a r. h i r s u t us ( H a r l an et a l ., 1970). Giant b e r m u d a g r a s s, C. d a c t y l on v a r. A r i d u s, is diploid a nd a p r o b a b le p r o g e n i t or of C. d a c t y l on v a r. d a c t y l o n. C r o s s es h a ve b e en m a de a m o ng and b e t w e en a ll of t h e s e, with t he b e st k n o wn b e i ng t he t r i p l o id c u l t i v a rs f r om C. t r a ns v a a l e n s is x C. d a c t y l on v a r. d a c t y l o n; T i f f i n e, T i f g r e e n, T i f w ay and T i f d w a r f. U g a n d a g r a ss l a wn a re s e l e c t i o ns f r om C. d a c t y l on v a r. d a c t y l o n. D r. H u f f i ne at O k l a h o ma h as b e en t e s t i ng s o me of his c o l l e c t i o ns of C. p l e t us f or r o a d s i de t u rf and s e v e r al h a ve b e en g r o wn at T u c s o n. But t h e re h as b e en l i t t le p r o g r e ss w i th t h is t a x o n, t he chief o b j e c- t i on being t he v e ry h a i ry l e a v es of m o st is a c u l t i v ar of C. t r a n s v a a l e n s i s, w h i le U3 and T i f- i n c o m- t y p e s. Gene f l ow b e t w e en d i s t i n ct t y p es of b e r m u d a g r a ss is r e s- t r i c t ed by s t e r i l i ty and i n f e r t i l i t y. w i th s e e ds of s o me c r o s s es ( H a r l an et a l ., 1969, 1970) and in I n h i b i t ed g e r m i n a t i on o c c u rs m a ny c a s es F^ p l a n ts t e nd to be v e g e t a t i ve and h e ad p o o r l y. T he c a u s es of s t e r i l i ty a p p e ar to be c h r o m o s o m a l, not in t he s e n se of m e i o t ic i r r e g u l a r i t y, but m o re c r y p t ic f a c t o r s, p o s s i b ly s h o rt i n v e r s i on s e g m e n ts or d e l e t i o n s. T r a n s l o c a t i on r a c es a re k n o wn in C. d a c t y l on v a r. d a c t y l o n. W i t h in C. d a c t y l on v a r. d a c- t y l on and in o t h er v a r i e t i e s, r e l a t i v e ly i n f e r t i le p l a n ts a re v e ry c o m m o n, e v en in c o m m e r c i al s e ed f i e l d s. T he t y p i c al v i g o r o u s, p r o m i s i ng s e l e c t i on f or t u r f g r a ss is a l m o st a l w a ys i n f e r t i l e, and s t r o n g ly v e g e t a t i v e, s o me t y p es n e v er h e a d i n g. W h a t e v er c a u s es of i n f e r t i l i t y, c r o s s es of f e r t i le x i n f e r t i le n o r m a l ly give i n f e r t i le and o f t en p o o r ly h e a d i ng p r o g e n i e s, a good f e a t u re f or t u r f, but u n f o r t u n a te f or s e l e c t i on t o w a rd s e e d ed t y p es ting t he good p o i n ts of t h e ir r e s p e c t i ve p a r e n t s. i n c o r p o r a- t r a d i t i o n al t he A l t h o u gh s e ed a n a l y s ts r e p o rt l i t t le d i f f i c u l ty with s e ed d o r- m a n cy in c o m m e r c i al l o t s, m u ch of t he s e ed o b t a i n ed f r om b r e e d i ng l i n es ( w h e t h er o p e n - p o l l i n a t i o n, s e l f e d, or c r o s s e d) t e n ds to be d o r m a nt and p r o g e n i es a re o f t en l i m i t ed f or t h is r e a s o n. 50% or m o re of t he s e ed o b t a i n ed f a i ls to p r o d u ce s e e d l i n g s. M a ny s e l fs a re q u i te v i g o r o us and t he e x t r e m e ly w e ak s e l f ed s e e d l i n gs f a m i l i ar in o t h er c r o s s - p o l l i n a t ed s p e c i es a re s e l d om s e e n. h a ps t he p o or r e c o m b i n a t i o ns t e nd to be in t he d o r m a nt s e e d s. T h is would be in a c c o rd with t he o b s e r v a t i o ns of H a r l an et a l ., ( 1969, 19 70) of i n c r e a s ed d o r m a n cy w i th s e e ds f r om c e r t a in c r o s s e s. P e r h a ps P e r- T h e re is e n o r m o us g e n e t ic v a r i a b i l i ty a v a i l a b le f or e x p l o r a- t i on in b e r m u d a g r a s s. To u se p e st p r o b l e ms as an e x a m p l e, p l a n ts v a r y i ng f r om s u s c e p t i b i l i ty to n e ar i m m u n i ty to b e r m u da m i t e, R h o d e s g r a ss s c a l e, R u t h 's s c a le and s p i d er m i t es h a ve b e en o b- s e r v e d. R e s i s t a n ce h as not y et b e en f o u nd in t u r f t y pe p l a n ts f or a n ew p e st ( so f ar c o n f i n ed to s e ed f i e l d s ), t he pink m e a ly bug w h i ch h as b e en p a r t i a l ly a c c o u n t a b le f or t he p r e s e nt high p r i ce of b e r m u d a g r a ss s e e d. Only in giant b e r m u d a g r a ss h a ve p l a n ts f r ee of t h is p e st b e en i s o l a t e d. A n o t h er A r i z o na p r o b l em is i r on c h l o r o s i s. S o me g e n o t y p es b e c o me c h l o r o t ic v e ry r e a d i ly and o t h e rs m a i n t a in an a c c e p t a b le g r e en c o l o r. O b v i o u s l y, t h e re a re e q u a l ly wide r a n g es in c o l o r, l e a f i n e s s, cold t o l e r a n c e, w i n t er h a r d i n e s s, e t c ., w h i ch a re of in t h is p o t e n t i al v a l u e. One of t he m o re p r o m i s i ng d e v e l o p m e n ts p r o g r am t y p es is t he d i s c o v e ry t h at t u rf q u a l i ty in s o me i n f e r t i le t e n ds to be d o m i n a nt in F^ 's b e t w e en t h e se p l a n ts and good s e ed s e t t i ng p l a n ts w h i ch p o s s e ss f ew o t h er v i r t u e s. U s i ng h i gh s e ed s e t t i ng p o t e n t i al as f e m a le p a r e n ts p r o v i d es an o b v i o us o p p o r t u n i ty f or p o t e n t i al c o m m e r c i al s e ed p r o d u c t i o n. p l a n ts with B e r m u d a g r a ss B r e e d i ng P r o g r am O b j e c t i v es of t he b r e e d i ng p r o g r am h a ve b e en to s e l e ct or d e v e l op high c o m b i n i ng c l o n es w h i ch w i ll p r o d u ce p r o f i t a b le y i e l ds of " h y b r i d" s e ed t h at will, in t u r n, p r o v i de c o n s u m e rs with s u- p e r i or t u r f g r a s s. T wo i m p o r t a nt f e a t u r es of b e r m u d a g r a ss m a ke t h is a p p r o a ch f e a s i b l e: (1) E a sy v e g e t a t i ve p r o p a g a t i o n, and (2) a h i gh l e v el of s e l f - s t e r i l i t y. S e l f ed s e ed s et is n o r m a l ly l e ss t h an .1% on a f l o r et b a s i s. S e l f - s t e r i le c l o n es p r o p a g a t ed v e g e t a t i v e ly in a l t e r n a te r o ws would p r o v i de s e ed f or p l a n t i ng in t he c o n s u m i ng a r e a. U s i ng only two c l o n es s h o u ld r e s u lt in a m u ch m o re u n i f o rm p r o d u ct t h an b r o a d er b a s ed c o m b i n a t i o ns or t h an a d v a n c ed s y n- t h e t ic g e n e r a t i o n s. N u r s e ry s t u d i es f r om m u t u al bag c r o s s i ng h a ve c o n f i r m ed t h i s. t he B e c a u se i n f e r t i l i ty and o t h er s e ed p r o d u c t i on p r o b l e ms a re f o r e m o st b o t t l e n e c ks to a s e e d ed v a r i e t y, a ll n ew m a t e r i a ls a re f i r st c h e c k ed f or s e ed p o t e n t i a l, hoping to r e t a in only t h o se with at l e a st s o me f e r t i l i ty f or t he b r e e d i ng p r o g r a m. S i n ce A r i z o na s e ed is p r o d u c ed p r i m a r i ly f or t he S o u t h e a s t e rn United S t a t e s, s e l e c t i o ns w i th d e m o n s t r a t ed v a l ue t h e re h a ve b e en of g r e a t er i n t e r e st t h an g e rm p l a sm f r om old of l o c al s e l e c t i o ns h a ve g i v en t u rf no d i f f e r e nt f r om c e r t i f i ed s e e d. U l t i m a t e l y, a body of s o u r ce m a t e r i a ls w i ll be a c c u m u l a t ed w h i ch w i ll c o m b i ne q u a l i ty p o t e n t i a ls w i th at l e a st r e a s o n a b le s e ed p r o d u c t i o n. T he g e n e t ic v a r i a b i l i ty f or m o st t r a i ts of v a l ue f r om a t u rf s t a n d p o i nt is a v a i l a b l e. S o me c r o s s es p r o d u ce high q u a l i ty p r o g e n y. S e ed s et f r om s u ch c r o s s es m u st be r a i s ed f r om 5% to 85%. S o me p l a n ts in c o m m e r c i al s e ed f i e l ds h a ve s e ed s e ts t h is h i gh or h i g h e r. V i g o r o us s e l e c t i o ns f r om t he s o u t h e a st g e n e r a l ly h a ve s e ed s e ts of 5% or l o w e r. R e s e a r ch is b e i ng done on s e ed p r o d u c t i on p r a c t i c e s, w h i ch m ay h e lp to e a se t h is p r o g l e m. G r o w e rs s t a n ds in A r i z o n a. C o m p o s i t es f or e x a m p l e, o f t en i r r i g a te t oo h e a v i ly f or b e st r e s u l t s. g e r m i n a t i on s t u d i es at T u c s on s o me y e a rs ago s u g g e s t ed h e a vy c o n c e n t r a t i o ns of p o l l en w e re n e c e s s a ry f or good s e ed set a nd p o l l en l o n g e v i ty w as l i m i t e d, m a k i ng e x t r e me i s o l a t i on d i s- t a n c es u n n e c e s s a r y. O u t c r o s s i ng s t u d i es w i th i s o l a t ed p l a n ts of giant b e r m u d a g r a ss d i s t a n c es f r om o t h er giant b e r m u d a g r a ss p l a n ts a re as l i t t le as 30 f e e t, t he diploid giant b e r m u d a g r a ss p l a n ts w i ll p r o d u ce up to 75% t r i p l o id o u t c r o s s es to t he t e t r a p l o id b e r m u d a g r a s s. It is a s s u m ed t h at t h e re is s i m i l ar d e p e n d e n ce u p on n e a r by p o l l en s o u r c es h as not b e en t e s t ed as yet. in b e r m u d a g r a s s. L a c k i ng good g e n e t ic m a r k e r s, in b e r m u d a g r a ss s e ed f i e l ds s u g g e st t h at w h en P o l l en t h at t h is R e f e r e n c es C i t ed H a r l a n, J. R ., and J. M. J. d e W e t. 1969. S o u r c es of v a r i a t i on in C y n o d on d a c t y l on ( L.) P e r s. C r op Sci. 9 : 7 7 4 - 7 7 8. H a r l a n, J. R ., J. M. J. d e W e t, and W. L. R i c h a r d s o n. 1969. H y- b r i d i z a t i on s t u d i es w i th s p e c i es of C y n o d on f r om E a st A f r i ca a nd M a l a g a s y. A m, J. Bot. 56:944-9 50. H a r l a n, J. R ., J. M. J. d e W e t, K. M. R a w a l, M. R. F e l d e r, and W. L. R i c h a r d s o n, 19 70. C y t o g e n e t ic s t u d i es in C y n o d on L. C. R i c h. ( G r a m i n e a e) C r op Sci. 10:288-291. K n e e b o n e, W. R. 1966. B e r m u d a g r a ss - - w o r l d l y, wily, w o n d e r f ul w e e d. E c o n. Bot. 2 0 : 9 4 - 9 7. I M P R O V E M E NT OF K E N T U C KY B L U E G R A SS F OR T HE T R A N S I T I ON Z O NE L i n c o ln H. T a y l or D e p a r t m e nt of A g r o n o my V i r g i n ia P o l y t e c h n ic I n s t i t u te & S t a te U n i v e r s i ty B l a c k s b u r g, V i r g i n ia 24061 I n t r o d u c t i on In t he E a s t e rn U n i t ed S t a t es it is d i f f i c u lt to g r ow s a t i s f a c t o ry t u r f g r a ss in an a r ea of t r a n s i t i on b e t w e en c o o l - h u m id and w a r m- h u m id c o n d i t i o n s. In t h is z o n e, c o o l - s e a s on g r a s s es do p o o r ly b e- c a u se of r e l a t i v e ly high t e m p e r a t u r e s, high h u m i d i ty and d r o u g h t, w h i le w a r m - s e a s on g r a s s es h a ve a long d o r m a nt s e a s on and m ay be s u b j e ct to w i n t e r k i ll (Holt, 19 69). No t u r f g r a ss s p e c i es is w e ll a d a p t ed in t h is t r a n s i t i on zone. T he p r e d o m i n a nt t u r f g r a s s es u s ed a re K e n t u c ky b l u e g r a ss v a r i e t i es and m i x t u r es of K e n t u c ky b l u e g r a ss a nd f i n e - l e a v ed f e s c u e s. B e r m u d a g r a ss and z o y s ia p e r f o rm w e ll as s u m m er w i n t er d o r m a n c y. T a ll f e s c ue p r o v i d es an a c c e p t a b le c o a r s e - l e a v ed l a wn in a r e as w h e re K e n t u c ky b l u e g r a ss is b e t t er a d a p t ed t h an K e n t u c ky b l u e g r a ss to a r e as of h a rd u s e, s t e ep s l o p e s, and s u n ny e x p o s u r es a t ., 19 69). t u rf but a re d i s l i k ed b e c a u se of t h e ir in t he t r a n s i t i on z o ne ( J u s ka et long p e r i od of is d i f f i c u lt to m a i n t a in and In V i r g i n i a, K e n t u c ky b l u e g r a ss and o t h er c o o l - s e a s on s p e c i es a re a d a p t ed to t he N o r t h e rn P i e d m o nt and a r e as c l o se to or w e st of t he B l ue R i d ge M o u n t a i n s. M i x t u r es of M e r i on K e n t u c ky b l u e g r a ss w i th K e n b l ue or South D a k o ta C e r t i f i ed ( w i th s o me r ed f e s c ue a d d ed f or s h a d ed a r e a s) a re t he p r e d o m i n a nt t u r f g r a s s es a nd u s ed ( S h o u l d e rs et a l ., 19 69). E a s t e rn V i r g i n ia and t he S o u t h e rn P i e d m o nt (with about l /2 of t he a r ea of t he s t a te and p e r h a ps 2 /3 of t he l a w ns and o t h er t u r f g r a s s) is in t he t r a n s i t i on zone. H e re K e n t u c ky b l u e g r a ss w i th r ed f e s c ue or K e n t u c ky 31 f e s c ue is r e c o m- m e n d ed f or s h a d ed l a w ns w i th b e r m u d a g r a s s, z o y s ia or K e n t u c ky 31 t a ll f e s c ue r e c o m m e n d ed f or o p en s u n l i g ht and p a r t i al s h a d e. A ll of t h e se s p e c i es h a ve p r o b l e ms w i th e s t a b l i s h m e n t, m a n a g e m e nt and a d e q u a te p e r f o r m a n ce in t he t r a n s i t i on z o n e. r e c o m m e n d ed T e m p e r a t u re F a c t or T e m p e r a t u re is a c r i t i c al f a c t or in a d a p t a t i on of t u r f g r a ss ( T h e se a re c a l c u l a t ed a c c o r d i ng to a s u g g e s t i on s p e c i e s. E s t i m a t es of e f f e c t i ve d ay and night t e m p e r a t u r es w h i ch s e v e r al g r a s s es a re w e ll a d a p t ed h a ve b e en m a de by M a d- i s on ( 1971). m a de by Went who t a k es t he d i f f e r e n ce b e t w e en t he m e an m i n i m um a nd m e an m a x i m um t e m p e r a t u re and d i v i d es it by f o u r. T h is f i g u re is a d d ed to t he m e an t e m p e r a t u re to get an e f f e c t i ve d ay t e m p e r a t u re and s u b t r a c t ed f r om t he m e an to get an e f f e c t i ve night t e m p e r a t u r e .) T he a p p r o x i m a te s u m m er t e m p e r a t u re r a n g es f or s e v e r a l t u r f g r a s s es a r e: to S p e c i es R ed f e s c ue K e n t u c ky b l u e g r a ss B e r m u d a g r a ss Z o y s ia E f f e c t i ve night t e m p e r a t u r e, °F E f f e c t i ve d ay t e m p e r a t u r e, °F 55 - 65 58 - 68 70 - 82 75 - 85 61 - 71 69 - 79 80 - 92 84 - 94 f r om F i g u re 4.6 of M a d i s on T he a r ea b e t w e en K e n t u c ky b l u e g r a ss and b e r m u d a g r a ss ( w h i ch do not o v e r l a p) is c o n s i d e r ed a t r a n s i t i on z o ne with no t u r f g r a ss s p e c i es w e ll a d a p t e d. E f f e c t i ve d ay and night t u r es f or s e v e r al l o c a t i o ns in V i r g i n ia w e re d e t e r m i n ed f r om l o n g - t i me s u m m er t e m p e r a t u re d a t a. B l a c k s b u rg and a r e as w e st of t he B l ue R i d ge w e re f o u nd to be w e ll w i t h in t he a r ea of K e n t u c ky b l u e g r a ss a d a p t a t i o n, R o a n o ke and L y n c h b u rg on t he f r i n ge of t h is a r e a, R i c h m o nd and t he a r ea n e ar W a s h i n g t o n, D . C. in t he t r a n s i- t i on z o n e, and N o r f o lk on t he e d ge of t he a r ea of b e r m u d a g r a ss a d a p t a t i o n. t e m p e r a- C l i m a t ic F a c t or In a d d i t i on to a ir t e m p e r a t u r e, W a rd (19 69) in h is c o n s i d e r a- t i on of c l i m a t ic f a c t o rs t h at a f f e ct t u r f g r a ss p e r f o r m a n c e, m e n t i o ns s o il t e m p e r a t u r e, m o i s t u r e, h u m i d i t y, r a d i a nt e n e r g y, wind and m i c r o c l i m a t e. M o i s t u re f r om p r e c i p i t a t i on d u r i ng t he g r o w i ng s e a s on is i m p o r t a nt to t he p e r f o r m a n ce of t u r f g r a ss in V i r g i n ia w h e re e a ch y e ar t h e re is a 10% p r o b a b i l i ty of 12 or m o re c o n s e c u- t i ve d r o u g ht d a y s. D r o u g h ts a re e v en m o re c o m m on in t he e rn P i e d m o nt and E a s t e rn V i r g i n ia t h an t he s t a te as a w h o le ( V a n B a v el and L i l l a r d, 19 57). M i c r o c l i m a t e, d e f i n ed as c l i m a te n e ar t he s o il s u r f a c e, is i n f l u e n c ed by s l o p e, s h a d e, m u l c h e s, and o t h er f a c t o r s. t h at o v er a t h r ee y e ar p e r i od in m i d s u m m er t he a v e r a ge a m o u nt of a K e n t u c ky b l u e g r a ss s t a nd a f f e c t ed by H e l m i n t h o s p o r um v a g a ns on a s o u t h - f a c i ng s l o pe w as 55% w h i le t wo n e a r by n o r t h- f a c i ng s l o p es a v e r a g ed 6% and 8% a f f e c t e d. In W e st V i r g i n i a, B e n n e tt and E l l i o tt ( 19 72) r e p o r t ed S o u t h- I m p r o v e m e nt of K e n t u c ky b l u e g r a ss f or t he t r a n s i t i on zone is f u r t h e r ed by t he g r e at d i v e r s i ty p r e s e nt w i t h in t he s p e c i es P oa p r a t e n s is L. and t he a p o m i c t ic c h a r a c t er w h i ch m a k es it p o s s i b le to u t i l i ze s u p e r i or c h a n ce or p l a n n ed h y b r id g e n o t y p e s. E x t e n s i ve c l o n al c o l l e c t i o n s, e s p e c i a l ly f r om s i t es not c o n s i d e r ed f a v o r a b le f or b l u e g r a s s, w i ll be e v a l u a t ed f or t u rf q u a l i t y. A c l o se c o r r e- l a t i on w as f o u nd by F u nk et al. ( 19 67) f or t u rf q u a l i ty of c l o n al a nd m a s s - s e e d ed p l o t s. S o me s e l e c t i o ns f r om old t u rf a r e as m ay be u s e f ul d i r e c t l y. O t h er s e l e c t i o ns w i th c h a r a c t e r i s t i cs to be of v a l ue f or p e r f o r m a n ce in t he t r a n s i t i on z o ne w i ll be u s ed as p a r e n ts to t h at p r o p o s ed by F u nk and H an ( 19 67) and r e p o r t ed by P e p in and F u nk (19 71) to be s u c c e s s f ul in p r o d u c i ng p r o m i s i ng a p o m i c t ic h y- b r i d s. in an i n t r a s p e c i f ic h y b r i d i z a t i on p r o g r am s i m i l ar t h at a p p e ar A d a p t a b i l i ty of K e n t u c ky B l u e g r a ss T h e re a re only l i m i t ed d a ta a v a i l a b le on c h a r a c t e r i s t i cs of K e n t u c ky b l u e g r a ss s t r a i ns w h i ch a re i m p o r t a nt to t h e ir p e r f o r- m a n ce u n d er s u m m er t e m p e r a t u re s t r e s s es in t he t r a n s i t i on z o n e. W i th t h r ee b l u e g r a ss s t r a i ns of d i s t i n c t ly d i f f e r e nt o r i g i ns g r o wn at f o ur t e m p e r a t u re r e g i m e s, Y o u n g n er and N u d ge (1968) f o u nd t h at M e r i o n, a s t r a in t h at o r i g i n a t ed in an a r ea of r e l a t i v e ly h i gh J u ly t e m p e r a t u r e s, a nd to p r o d u ce m o re t op g r o w th t h an F y l k i ng or N e w p o r t, w h i ch o r i g i n a t ed in c o o l er a r e a s. F i ve K e n t u c ky b l u e g r a s s es N u g g e t, K e n b l ue and t wo s e l e c t i o ns m a de n e ar N o r f o l k, V i r g i n ia w e re g r o wn at t h r ee t e m p e r a t u re r e g i m es and at high and l ow n i t r o- g en l e v e ls by W a t s c h ke et al. (1970). T h ey c o n c l u d ed t h at b l u e g r a s s es t e n d ed to h a ve t he h i g h e st c a r b o h y d r a te - P e n n s t a r, r e s e r v es - t e m p e r a- In a f u r t h er s t u d y, W a t s c h ke et al. o r i g i n a t i ng in w a rm r e g i o ns w e re b e t t er a d a p t ed to h i gh t u re s t r e ss b e c a u se of i n h e r e nt l o w er n i t r a te a b s o r p t i on and h i g h er c a r b o h y d r a te l e v e l s. g r ew t en b l u e g r a ss s t r a i ns u n d er high t e m p e r a t u re s t r e ss f or f o ur w e e ks and r e c o r d ed f o l i ar y i e ld and c a r b o h y d r a t es and r a t es of p h o t o s y n t h e s is and r e s p i r a t i o n. B l u e g r a s s es w i th b e st g r o w th at h i gh t e m p e r a t u r es u s u a l ly w e re h i g h e st in c a r b o h y d r a t e s, had h i g h er p h o t o s y n t h e s is and l o w er r e s p i r a t i o n. T h ey s u g g e s t ed b l u e g r a s s es with g e n e t i c a l ly high p h o t o s y n t h e t ic e f f i c i e n cy be s e l e c t ed f or u se in a r e as of h i gh t e m p e r a t u re s t r e s s e s. (19 72) t h at No l a b o r a t o ry t e c h n i q ue p r e s e n t ly a v a i l a b le would a p p e ar of g r e at p r a c t i c al v a l ue f or s c r e e n i ng l a r ge q u a n t i t i es of K e n- t u c ky b l u e g r a ss b r e e d i ng m a t e r i a ls f or a d a p t a t i on to high t e m- p e r a t u re s t r e s s e s. S o me e v a l u a t i on f or c a r b o h y d r a te p h o t o s y n t h e t ic e f f i c i e n c y, b e i ng e v a l u a t ed as p o t e n t i al p a r e n ts But if we n e ed s t r a i ns t h at w i ll p e r f o rm u n d er high s t r e s s es we w i ll n e ed to e v a l u a te u n d er h i gh t e m p e r a t u re A m a j or p a rt of t h is e v a l u a t i on m i g ht m a ke u se of t he v a r i- a b le and v a r i o us s t r e s s es t h at n a t u re p r o v i d es so g e n e r o u s ly t he t r a n s i t i on zone. etc. m i g ht be d e s i r a b le f or s t r a i ns in h y b r i d i z a t i on p r o g r a m. l e v e l, t e m p e r a t u re in s t r e s s e s. R e f e r e n c es C i t ed B e n n e t t, O. L., and E. S. E l l i o t t. i n c i d e n ce on f i ve f o r a ge s p e c i es as a f f e c t ed by n o r th and s o u th f a c i ng s l o p e s. P l a nt D i s e a se R e p t r. 19 72. P l a nt d i s e a se 5 6 : 3 7 1 - 3 7 5. F u n k, C. R ., R. E. E n g e l, and P. M. H a l i s k y. 1967. C l o n al e v a l u a- t i on of s e l e c t i o ns of K e n t u c ky b l u e g r a s s, P oa p r a t e n s is L. p. 5 5 - 67 J e r s ey A g r. E x p. Sta. Bull. In 1967 R e p o rt of T u r f g r a ss R e s e a r c h. N ew 818. F u n k, C. R., and S. J. Han. 19 67. R e c u r r e nt i n t r a s p e c i f ic h y b r i d i- a p r o p o s ed m e t h od of b r e e d i ng K e n t u c ky b l u e g r a s s, z a t i o n: P oa p r a t e n s is L. p. 3-14 R e s e a r c h. N ew J e r s ey A g r. E x p. Sta. Bull. 818 In_ 19 67 R e p o rt of T u r f g r a ss H o l t, E. C. 19 69. T u r f g r a s s es u n d er w a r m, h u m id c o n d i t i o n s. In T u r f g r a ss S c i e n c e. A g r o n o my M o n o g r a ph 14. A m e r i c an S o- c i e ty of A g r o n o m y. M a d i s o n, W i s e. J u s k a, F. V., J. F. C o r n m a n, and A. H. Hovin. 1969. T u r f g r a s s es u n d er cool, h u m id c o n d i t i o n s. In T u r f g r a ss S c i e n c e. A g r o n o my M o n o g r a ph 14. A m e r i c an S o c i e ty of A g r o n o m y. M a d i s i o n, W i s e. M a d i s o n, J. H. 1971. P r i n c i p l es of T u r f g r a ss C u l t u r e. V an N o s- t r a nd R e i n h o ld Co. N ew Y o r k. P e p i n, G. W., and C. R. F u n k. 19 71. I n t r a s p e c i f ic h y b r i d i z a t i on as a m e t h od of b r e e d i ng K e n t u c ky b l u e g r a ss ( P oa p r a t e n s is L.) f or t u r f. C r op Sci. 11:445-448. S h o u l d e r s, J. F ., R. E. S c h m i d t, R. Es B l a s e r, and A. S. B e e c h e r. 1969. T u r f g r a ss guide f or l a w ns and o t h er t u rf a r e a s. P u b l i c a t i on 311. C o o p e r a t i ve E x t e n s i on S e r v i c e. V i r g i n ia P o l y t e c h n ic and S t a te U n i v e r s i t y. I n s t i t u te V an B a v e l, C. H. M., and J. H. L i l l a r d. 19 57. A g r i c u l t u r al d r o u g ht in V i r g i n i a. T e c h. Bull. 128. Va. A g r. E x p. Sta. W a r d, C. Y. 1969. C l i m a te and a d a p t a t i o n. In T u r f g r a ss S c i e n c e. A g r o n o my M o n o g r a ph 14. A m e r i c an S o c i e ty of A g r o n o m y. M a d i s o n, W i s e. W a t s c h k e, T. L., R. E. S c h m i d t, and R. E. B l a s e r. 1970. R e s p o n s es of s o me K e n t u c ky b l u e g r a s s es to h i gh t e m p e r a t u re and n i t r o g en f e r t i l i t y. C r op Sci. 1 0 : 3 7 2 - 3 7 6. W a t s c h k e, T. L., R. E. S c h m i d t, E. W. C a r s o n, and R. E. B l a s e r. 1972. S o me m e t a b o l ic p h e n o m e na of K e n t u c ky b l u e g r a ss u n d er high t e m p e r a t u r e. C r op Sci. 12:87-90. Y o u n g n e r, V. B., and F. J. N u d g e. 19 68. G r o w th and c a r b o h y d r a te s t o r a ge of t h r ee P oa p r a t e n s is L. s t r a i ns as i n f l u e n c ed by t e m p e r a t u r e. C r op Sci. 8 : 4 5 5 - 4 5 7. T U R F G R A SS B R E E D I NG IN F L O R I DA A l l en E. D u d e ck U n i v e r s i ty of F l o r i da A g r i c u l t u r al R e s e a r ch C e n t er F o rt L a u d e r d a l e, F l o r i da 33314 I n t r o d u c t i on A u g u s t i n e g r a s s, b a h i a g r a ss and b e r- B r e e d i ng w o rk on w a rm s e a s on t u r f g r a s s es is b e i ng c o n d u c- E m- t ed at t he A g r i c u l t u r al R e s e a r ch C e n t er at F o rt L a u d e r d a l e. p h a s is is being p l a c ed on St. m u d a g r a s s, a l t h o u gh a s e l e c t i on p r o g r am with z o y s i a g r a ss and c e n t i p e d e g r a ss g r am at F o rt L a u d e r d a le a re E. O. B u r t, t u rf t e c h n o l o g i s t; R. E. M c C o y, p a t h o l o g i s t; J. A. R e i n e r t, e n t o m o l o g i s t; S. R. J o h n s o n, n e m a t o l o g i st at B e l le G l a d e; and G. C. H o r n, t u rf t e c h n o l o g i st at G a i n e s v i l l e. is a l so b e i ng c a r r i ed out. C o o p e r a t i ng in t h is p r o- St. A u g u s t i n e g r a ss T he p r i m a ry o b j e c t i ve in St. A u g u s t i n e g r a ss is to d e v e l op t y p es w h i ch a re r e s i s t a nt to St. A u g u s t i n e g r a ss d e c l i ne v i r us (SAD), c h i n ch b u g s, b r o wn p a t ch and r o ot knot n e m a t o d e s. D w a r f, l o w - g r o w i ng t y p es w i th d a r k - g r e en c o l or and m i n i m al t h a t c h i ng t e n d e n c i es a re a l so d e s i r e d. i m p r o v e m e nt In c o o p e r a t i on with T e x as A&M U n i v e r s i t y, F l o r i da is r e- l e a s i ng a n ew St. A u g u s t i n e g r a ss to be n a m ed " F l o r a t a m ." T h is v i g o r o us v a r i e ty is r e s i s t a nt to SAD and downy m i l d e w, and d o es not s h ow zinc and i r on d e f i c i e n c i es u n d er T e x as c o n d i t i o n s. L a b o r a t o ry and f i e ld t e c h n i q u es h a ve b e en d e v e l o p ed f or q u i ck s c r e e n i ng of St. A u g u s t i n e g r a s s es f or R h i z o c t o n ia s o l a ni a nd c h i n ch bug t o l e r a n c e. P h o t o - p e r i od s t u d i es a re u n d e r w ay to m a n i p u l a te f l o w e r i ng f or c o n t r o l l ed c r o s s i ng in t he g r e e n h o u s e. B a h i a g r a ss T he p r i m a ry o b j e c t i ve in b a h i a g r a ss i m p r o v e m e nt is to d e- v e l op t y p es w h i ch a re d e t e r m i n a te in t h e ir f l o w e r i n g. F l o r i d a, b a h i a g r a ss f l o w e rs c o n t i n u o u s ly f r om M ay to N o v e m b er a nd r e q u i r es e x c e s s i ve m o w i ng to p r o d u ce an a e s t h e t i c a l ly d e- s i r a b le In South t u r f. B e r m u d a g r a ss i m p r o v e m e nt T he p r i m a ry o b j e c t i ve in b e r m u d a g r a ss is to d e v e l op t y p es w h i ch a re r e s i s t e nt to sting and l a n ce n e m a t o d e s, b e r m u d a g r a ss m i te and leaf spot ( H e l m i n t h o s p o r i um spp.). S e- l e c t i o ns f or v i g o r o us t y p es u n d er m i n i m um f e r t i l i z a t i on w e re m a d e. T e m p e r a t u re t y p es w h i ch w i ll g r ow y e a r - r o u nd in South F l o r i da a re d e s i r e d. A r a p id s c r e e n i ng t e c h n i q ue s t a r ch a c c u m u l a t i on u n d er c o ol n i g h ts is b e i ng u s ed to i d e n t i fy p r o m i s i ng t y p e s. involving P r i m a ry a c t i v i t i es to d a te i n v o l v ed c a t e g o r i z i ng a p p r o x i m a t e ly 1,300 c l o n e s. Upon c o m p l e t i o n, s i n g le or p o l y c r o ss h y b r i d i z a t i on w i ll be a t t e m p t e d.