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I I I I 0. I I I. . . . I .I. . I , I I II . I 1 I I I I Q.I . .J I . . .. «I... . £99. I. III I o! IIIII . 0.0. I I III- II... I.\. I . I . .. . . . .. I I. I. I I #9.; ‘1... cl. .. ‘ a. I. . 9 III4 . H \II .0 I .. . I I .III 1a....o o ..I III... .3- . - I . . . .. _ 1...... . .. I a... . . .31 a? $3.965... xi. .4. _ ¢ ., fl . I I . I I0. I I . I 0. I o . . . .. I II I «Is 0.... \ .o.r' .. .m. ' I IIII. III... I! I II... .I .5; .0 y. Y . .I . . . 31.. . .« P1. 0 I ‘I. ll I III“ II I- I ll III I ' II 1'. II I'll-r I . .l- I 0 _. .- PLACE IN RETURN BOX to remove this checkout from your record. TO AVOID FINES return on or before date due. MAY BE RECALLED with earlier due date if requested. DATE DUE DATE DUE DATE DUE 5I08 K:IProlecc&PreleIRC/DateDue.indd ABSTRACT SEED TREATMENTS LEADING TO INCREASED RATES OF SHOOT EMERGENCE OF FOUR COOL-SEASON TURFGRASSES BY Richard B. Anda The object of this study was to find a seed treatment which would increase the rate of shoot emergence and thus reduce losses during establishment due to erosion, suboptimal growth conditions, or weed competition. Soaking and Drying Results Four cool season turfgrasses seeds, Manhattan perennial ryegrass (Lolium perenne L.), Merion Kentucky bluegrass (Poa pratensis L.), Pennlawn red fescue (Festuca rubra L.), and Penncross creeping bentgrass (Agrostis palustris Huds.) were soaked at temperatures of 5, 10, 15, 25, and 15-25 C, with soaking times of 6, 12, 24, 48, and 168 hours. Soaking solutions of distilled water and polyethylene glycol were also compared. Drying treatments were either air dried at 23.9 C for one week or oven dried at 45 C for 18 hours. Petri dish studies in the growth chamber were used for the majority of the studies, along with one field experiment. The Pen for 168 hours 10 C soaking f soaking treatm soaking-drying by air drying emergence. 0: 5C for 48 hor soaking or dry time of the 24 increased the bluegrass. A} oven drying f, and Pennlawn, t0 germinatim Thus 5. bluegrass Soa' Of emergence be adaptable grass areas, for all tur fg Richard B. Anda The Penncross soaking treatments of 5, 15, and 15-25 C for 168 hours along with the soaking-drying treatment of 10 c soaking for 48 hours followed by air drying; and the soaking treatment for Merion of 25 C for 168 hours and the soaking-drying treatment of 10 C soaking for 48 hours followed by air drying were effective in increasing the rate of shoot 'emergence. Only soaking treatments of 25 C for 6 hours and 5 C for 48 hours were found effective for Pennlawn, while no soaking or drying treatments were effective with Manhattan. None of the 24 polyethylene glycol seed soaking treatments increased the rate of shoot emergence of Merion Kentucky bluegrass. Also, air drying was significantly better than oven drying for Merion and Penncross, where as with Manhattan and Pennlawn, oven drying after soaking was more detrimental to germination than air drying. Thus seed soaking, and in the case of Merion Kentucky bluegrass soaking followed by drying, will increase the rates of emergence of Merion, Pennlawn, and Penncross, and should be adaptable to establishment by hydroseeding of most turf- grass areas. No one treatment was found to be effective for all turfgrasses, since each reacted differently to soaking time and temperature. Kentucky Bluegrass Treatment Effects During and After Anthesis Since the growth of perennial ryegrass (Lolium perenne L.) was found to be increased by foliar applications of simazine and atrazine, and also since foliar applications of urea on wheat (Triticum aestivum L.) increased yield and protein conteni ffect such fol germination of treatments. K 'Herion' and ' of simazine ar one and three was at anthes beinc; seven 2 treatments are agent. The r “flirtatious Sign'lficantj Seed germin establiShme cor Richard B. Anda protein content, this study was undertaken to determine what effect such foliar applications would have on the rate of germination of seeds formed under the influence of these treatments. Kentucky bluegrasses (Poa pratensis L.) 'Merion' and 'Kenblue' were treated with spray applications of simazine and urea during and after anthesis with both one and three application treatments. The first application was at anthesis, with the second and third applications being seven and fourteen days after anthesis. Identical treatments were made with and without using a wetting agent. The results indicate that none of the treatment combinations with urea, simazine, and a wetting agent were significantly better than the control in terms of percent seed germination at 5, 10, 15, 20, and 30 days after establishment, according to Dunnett's test for two-sided comparisons at the 5% level. in Par' Depe SEED TREATMENTS LEADING To INCREASED RATES OF SHOOT EMERGENCE OF FOUR COOL-SEASON TURFGRASSES BY Richard Brian Anda A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Department of CrOp and Soil Sciences 1975 The a persons wit? have been as Mr. Bailey of r; 5&1"! seed 1 Dr. constructi most “mpg! deVelop’ Dr and Dr. J stud i e S E M E encour ac ACKNOWLEDGEMENTS The author expresses his appreciation to the following persons without whose presence this study would not otherwise have been as complete, meaningful, and rewarding: Mr. Arden Jacklin of Jacklin Seed Company and Mr. Dick Bailey of Turfgrass Seed Incorporated who provided the neces- sary seed for the study. Dr. James B. Beard for his guidance, encouragement, constructive criticism in reviewing the manuscripts, and most importantly the freedom he provided me to determine, develop, and carry out this thesis research. Dr. Larry Copeland, Dr. Ken Payne, Dr. Paul Rieke, and Dr. Joe Vargas, Jr. who assisted throughout my graduate studies and served on my guidance committee. My parents who have given their total support and encouragement throughout my education. Diane for her research assistance, enthusiasm, encouragement, sacrifice, and love. ii ACKNOWLEDGEMEN LIST OF TABLES LIST 0? Appm INTRODUCTION . SOAKING AND D] EMERGENCE 1 TURFGRASSE: Abstrac Introdu Literat Materia REBSultE REfereI TABLE OF CONTENTS ACKNOWLEDGEMENTS . . . . . . LIST OF TABLES . . . . . . . LIST OF APPENDIX TABLES . . INTRODUCTION . . . . . . . . SOAKING AND DRYING TREATMENTS USED TO INCREASE EMERGENCE RATES OF FOUR COOL-SEASON TURFGRASSES . . . . . . . Abstract . . . . . . . Introduction . . . . . Literature Review . . Materials and Methods Results and Discussion References . . . . . . EFFECT OF FOLIAR APPLICATIONS OF DURING AND AFTER ANTHESIS ON SEED GERMINATION OF POA PRATENSIS L. 'MERION' Abstract . . . . . . . Introduction . . . . . Materials and Methods Results and Discussion References . . . . . . CONCLUSIONS . . . . . . . . APPENDIX C O O O I O O O O 0 iii UREA AND SIMAZINE AND 'KENBLUE' . . Page ii iv vi \IU'IUJLU 19 25 34 49 49 50 52 55 59 60 63 Table Bentore creepii which I the un* observe Dunnet' Bent r; creepi; treatm. differ. a sPec. leVel . compar FESCue fescue Signif COntrO at the two‘si FeSCue feSCUE Which the HI ObSEr\ Duhnet Pe “ha: treatI diffe: cOmPa: Table 1. LIST OF TABLES Bentgrass Soaking Treatments. Penncross creeping bentgrass seed soaking treatments which were significantly different from the untreated control, at a specific observation date, at the 5% level using Dunnett's test for two-sided comparisons . Bentgrass Drying Treatments. Penncross creeping’bentgrass seed soaking-drying treatments which were significantly different from the untreated control, at a specific observation date, at the 5% level using Dunnett's test for two-sided comparisons . . . . . . . . . . . . . . Eescue Soakinngreatments. Pennlawn red fescue seed soaking treatments which were significantly different from the untreated control, at a specific observation date, at the 5% level using Dunnett's test for two-sided comparisons . . . . . . . . . Fescue Drying Treatments. Pennlawn red fescue seed soaking-drying treatments which were significantly different from the untreated control, at a specific observation date, at the 5% level using Dunnett's test for two-sided comparisons Perennial Ryegrass Soaking Treatments. Manhattan perennial ryegrass seed soaking treatments which were significantly different from the untreated control, at a specific observation date, at the 5% level using Dunnett's test for two-Sided comparisons . . . . . . . . . . . . . . iv Page 39 40 41 42 43 Table 10. ll. Perenr Manhat drying differ specif using compar Kentuc Merlor treatn differ a spec level compaz Kentuc Merior drying differ Specii using Compax The e1 Seedlj DIUEg] per dr Table 6. 10. 11. Perennial Ryegrass Drying Treatments. Manhattan perennial ryegrass seed soaking- drying treatments which were significantly different from the untreated control, at a specific observation date, at the 5% level using Dunnett's test for two-Sided comparisons . . . . . . . . . . . . . . . Kentucky Bluegrass Soaking Treatments. Merion Kentucky bluegrass seed soaking treatments which were significantly different from the untreated control, at a specific observation date, at the 5% level using Dunnett's test for two—sided comparisons . . . . . . . . . . . . . . . Kentucky Bluegrass Drying Treatments. Merion Kentucky bluegrass seed soaking- drying treatments which were significantly different from the untreated control, at a specific observation date, at the 5% level using Dunnett's test for two-sided comparisons . . . . . . . . . . . . . . . The effect of 12 seed treatments on seedling establishment of Merion Kentucky bluegrass in the field, counted in shoots per dm2 and as percent cover ratings . . . Effects of foliar applications of urea and simazine on the germination rate of Merion Kentucky bluegrass . . . . . . . . Effects of foliar applications of urea and simazine on the germination rate of Kenblue Kentucky bluegrass . . . . . . . . Page 44 45 46 47 57 58 Table 1. 10. LIST OF APPENDIX TABLES Page Penncross creeping bentgrass soaking treatment means for seed germination . . . . . . . 63 Penncross creeping bentgrass soaking- drying treatment means for seed germination . . . . . . . . . . . . . . . . . . . 64 Pennlawn red fescue soaking treatment means for seed germination . . . . . . . . . . . . 65 Pennlawn red fescue soaking-drying treatment means for seed germination . . . . . . . 66 Manhattan perennial ryegrass soaking treatment means for seed germination . . . . . . . 67 Manhattan perennial ryegrass soaking- drying treatment means for seed germination O I O O O O O O O O I O O O O O O O O 68 Merion Kentucky bluegrass soaking treatment means for seed germination . . . . . . . 69 Merion Kentucky bluegrass soaking- drying treatment means for seed germination O I O O O I O O O O O O O O O O O O O 70 Temperature and rainfall data from September 20 to October 15, 1974, in East Lansing, Michigan, as recorded by the U.S. Dept. of Commerce Climatological Data . . . . . . . . . . . . . . . . . . . . . . . 71 Polyethylene glycol seed soaking treatment germination means . . . . . . . . . . . 72 vi INTRODUCTION Establishment of some cool—season turfgrasses by seeding often includes problems with soil erosion and weed infestation before the turfgrass seed germinates and fully develops. Kentucky bluegrasses (Poa pratensis L.) are noted for their relatively slow germination rate compared to most other cool season turfgrasses (Parks and Henderlong - 1967). Attempts have been made at finding a method of reducing the time of Shoot emergence of many seed species through seed soaking, soaking followed by drying, and soaking with chemical stimulants, which have produced positive, negative, and no effect results. Yet, little research has been done with perennial cool-season turfgrasses (Frischknecht - 1959). In 1974, nearly 100 million pounds of cool-season turfgrass seed was produced and sold for establishment of permanent cover. Of that 100 million pounds, 57.3 million pounds of Kentucky bluegrasses (personal communication with Dr. R.W. Schery, The Lawn Institute). Therefore, since some cool-season turfgrasses require more than fifteen days to germinate and such a large quantity of seed is produced each year, there is a definite need and justification for research leading to increased rates of germination of the four major cool-season turfgrasses. l The objectives of this study were to determine (1) the effects on four turfgrasses of seed soaking and soaking followed by drying and (2) the effect of foliar applications on the inflorescence of two turfgrasses, with urea and sub- herbicidal rates of simazine on the rate of shoot emergence. Each aspect was studied individually, and the results are presented in two sections. It was intended that the results from these investiga- tions would be useful in improving turfgrass seedling emergence and therefore reduce the need for mixing rapid germinating species with slower germinating species when seeding to prevent soil erosion, reduce weed competition during establishment, and reduce the need for sod. SOAKING AND DRYING TREATMENTS USED TO INCREASE EMERGENCE RATES OF FOUR COOL-SEASON TURFGRASSES ABSTRACT The object of this study was to find a seed soaking or combination soaking-drying treatment which would increase the rate of shoot emergence and thus reduce losses during establishment due to erosion, suboptimal growth conditions, or weed competitionx’ Four cool-season turfgrasses, Manhattan perennial ryegrass (Lolium perenne L.), Merion Kentucky bluegrass (Poa pratensis L.), Pennlawn red fescue (Festuca rubra L.), and Penncross creeping bentgrass (Agrostis palustris Huds.) were used in this study. Seeds were soaked at temperatures of 5, 10, 15, 25, 15-25 C, with soaking times of 6, 12, 24, 48, and 168 hours. Soaking solutions of distilled water and polyethylene glycol were also compared. Drying treatments were either air dried at 23.9 C for one week or oven dried at 45 C for 18 hours. Petri dish studies in a growth chamber were used for the majority of the studies, along with one field experiment. The Penncross soaking treatments of 5, 15, and 15-25 C for 168 hours along with the soaking-drying treatment of 10 C soaking for 48 hours followed by air drying; and the soaking treatment for Merion of 25 C for 168 hours and the soaking- drying treatment of 10 C soaking for 48 hours followed by air drying were effective in increasing the rate of shoot emergence. .Only soaking treatments of 25 C for 6 hours and 5 C for 48 hours were found effective for Pennlawn, while no soaking or drying treatments were effective with Manhattan. None of the 24 polyethylene glycol seed soaking treatments increased the rate of shoot emergence of Merion Kentucky bluegrass. Also, air drying was significantly better than oven drying for Merion and Penncross, while with Manhattan and Pennlawn, oven drying after soaking was more-detrimental to germination than air drying. Thus hydroseeding, and in the case of Merion Kentucky bluegrass dry seeding, after a seed soaking treatment will increase the rates of emergence of Merion, Pennlawn, and Penncross, and would be adaptable to establishment of most turfgrass areas. No one treatment was found to be effective for all turfgrasses, since each reacted differently to soaking time and temperature. Kentur of 98) of ray erosic use 5‘ and c1 the t. Signi 100 m eStab Anoth 1974 COVeI INTRODUCTION Several cool season turfgrass species including Kentucky bluegrasses (Poa pratensis L.) have a slow rate of germination (Parks and Henderlong - 1967). The advantage of rapid germination rates is reduced proneness to soil erosion, the potential for weed infestation, and the need to use seeding mixtures containing rapidly germinating species and cultivars. Thus the advantages of being able to decrease the time between seeding and germination are extremely significant and worth investigating, especially since nearly 100 million pounds of cool-season turfgrasses were established last year specifically for permanent cover. Another 200 million pounds of turf type seed was sold in 1974 for winter cover and agricultural purposes (temporary cover). Kentucky bluegrass seed sold, excluding Merion, totaled 52.3 million pounds in 1974, while Merion was 5 million pounds. Red fescue produced in 1974 was 7.5 million pounds, Bentgrass produced was 9.2 million pounds, and all ryegrass produced was 220 million pounds with perennial ryegrass being 10-15 million pounds (personal communication with R.W. Schery, The Lawn Institute). Therefore, since some of the cool-season turfgrasses require more than fifteen days to germinate and such a large quantity of seed is produced each year, there is a definite need and justification for research leading to increased rates of germination of the four major cool-season turfgrasses. Many statements, both positive and negative, have been made regarding the effects of pregermination seed soaking, drying, and the use of chemical stimulants. Yet, little research has been done with perennial grasses, especially cool—season turfgrasses (Frischknecht - 1959). The objectives of this study were to determine: 1) if soaking turfgrass seeds in distilled water would stimulate the rate of seedling emergence; and, if so, what soaking temperature and time combinations are optimal, 2) if turfgrass seeds could be dried after being soaked and still retain an emergence rate significantly better than an untreated control, 3) if the rate of emergence of Merion Kentucky bluegrass could be improved by soaking treatments with polyethylene glycol, and 4) if, when Merion Kentucky bluegrass seed treatments selected from the three above mentioned treatment categories were established in the field, the rates of emergence would be significantly faster than an untreated control. . Qt! LITERATURE REVIEW Soaking With Water The effects of soaking seeds of different crop species in water before establishment has been reported by many investigators. Cases where soaked seeds germinated more rapidly than untreated seeds include: Kidd and West (1918b), Kidd and West (1918a), and Kidd and West (1919) with various crop seeds; Chippendale (1934) with some Gramineae spp.; Haight and Grabe (1972) with orchardgrass (Dactylis glomerata L.). Keller and Bleak (1968) found that crested wheatgrass (Agropyron cristatum L.) emerged more rapidly from the soil than the control when the seed was soaked for two to three days at room temperature; and Bleak and Keller (1969) also reported that crested wheatgrass which was soaked, but not submerged, emerged more rapidly than untreated seeds. The response was in direct proportion to the duration of seed soaking, with soaking also improving Shoot and root growth. Although the initial rate of emergence and growth was often better than the control, Kidd and West (1918b) with various crop seeds, Keller and Bleak (1969) with crested wheatgrass, and Haight and Grabe (1972) with orchardgrass found that the final total germination and growth were nearly equal. Frost and cold resistance have been found to increase in seedlings whose seeds had been soaked prior to establish- ment. These low temperature stress responses were reported by Kushnirenko (1957) with tomato (Lycopersicon Spp. Mill.) and corn (Zea mays L.), Kydrev (1959) with Mercury wheat (Triticum aestivum L.), Thomas and Christiansen (1971) with cotton (Gossypium spp. L.) and Cole and Wheeler (1974) with cotton. Genkel (1964) with several crops and Lyles and Fanning (1964) with grain sorghum (Sorghum vulgare Pers.) found that seeds germinating under drought stress which had been soaked before sowing emerged more rapidly than the control (Lyles) and produced drought tolerant plants (Genkel). On the other hand, some researchers state that soaking with water was ngt_beneficial to increased germination rates and development of a plant: a) Kidd and West (1918b) state that soaking seeds in large volumes of water reduces the beneficial effects, and may even be harmful to seeds, b) With orchardgrass Linehan and Mercer (1936) found no acceleration in laboratory germination after soaking seeds with water, c) Sorghum height, weight, and yield were lower under drought conditions with seeds which had been soaked and dried than with untreated seed (Evenari - 1964). Temperature of the soaking water appears critical, since 70 C soaking for 5 minutes reduced cotton seed germina- tion (Cole and Wheeler - 1974). Young, Kay, and Evans (1974) state that high temperature soaking must be with aeration and of very short duration. Soaking or Treating With Gibberellin, Hormones, and Other Compounds Gibberellic acid (GA), hormones, and various other extract compounds have been used as seed treatments at varying concentrations for the purpose of stimulating seed germination and seedling growth. Button (1959) found that GA hastened the germination of red fescue (Festuca rubra L.), domestic ryegrass (Lolium perenne L.), tall fescue (Festuca arundinacea Schreb.), and Kentucky bluegrass. Kentucky bluegrass, perennial ryegrass, and creeping red fescue each reacted with increased germination rates and leaf growth when Behrendt (1960) applied specific amounts of GA to soaking solutions for each species. Cole and Wheeler (1974) reduced chilling injury to cotton plants by soaking the seed in either GA or cyclic AMP. Young et a1 (1974) worked with bermudagrass (Cynodon dactylon L.) and reported that adding GA3 to the soaking solution enhanced the rate of germination and total germination, and than when kinetin was added to the GA3 in the soaking solution, seedling size increased. Seaweed extract used at 0.5% and 1.0% in a soaking solution for creeping red fescue increased the speed of seedling emergence (Button and Noyes - 1964). Since a water control was inferior to the seaweed treatment, it appears that the seaweed extract was effective. Nonstimulatory effects with GA have been reported by Andersen (1957) with Merion Kentucky bluegrass and by Haight and Grabe (1972) with orchardgrass. Both reported that GA 10 and water were of equal value in stimulating seed germination. Porter (1950) found that the following compounds had minimal response or were detrimental to dormant and nondormant seed species germination: ascorbic acid, colchicine, Hormodin "A", indoleacetic acid, indolebutyric acid, indol-3- acetic acid, K-a-naphthalene-acetate, lacto flavin, levulinic acid, naphthalene acetamide, naphthalene acetic acid, thiourea, vitamin B or 13 commercial hormone dusts. 1’ Soaking_with Dilute Nutrient Solutions or Fungicide Nelson (1927) stated that the Speed of germination and total number of seeds germinating in Pca_spp. were increased by presoaking the seed in a potassium nitrate (KNO3) solution. Andersen (1935) found that Canada bluegrass (Egg compressa L.) germination was not affected by nitrate solution when the seeds were in soil, but the percent germination increased substantially when seeds were given nitrate solution on filter paper. Bluegrass and tall fescue seedling survival was reported improved with potassium chloride but not with soaking in urea (Daniel and Goetze - 1957). With Cougar and Newport Kentucky bluegrass seed, Maguire and Steen (1971) found that 0.1% - 0.2% KNO3 soaking solution was more effective than water in increasing the germination rate, as well as increasing respiration rates. Martyanova (1960) reported that soaking barley seeds (Hordeum vulgare L.) with boric acid produced the best results in terms of emergence rates when compared to low temperature hardened and control treatments. 11 Others feel that dilute nutrient solutions and fungicides have no positive effect on seed germination. Tyler, Murphy, and MacDonald (1956) found that, except where smut was present, fungicide treatment of seed had little value in terms of increased stand or yield. Daniel and Goetze (1957) and Daniel (1958) found that among five fertilizers (potassium chloride, ammonium nitrate, nitrogen plus phosphorus, superphOSphate, or urea), none enhanced germination or seedling growth over water soaked seeds. Daniel (1958) also reported that potassium salts are less toxic to seeds during soaking than are ammonia salts. Finally, Dickens and Moore (1974) found that KNO3 did not affect the rate of emergence or the total number of seeds germinated in cogongrass (Imperata cylindrica L.). Drying After Soaking Treatments In order to facilitate seed planting, soaked seeds are often dried before being sown. The effects of the drying methods on the germination advantage derived from soaking are both positive and negative. Kreyger (1963) classified grass seeds as quick drying. He states that when drying by the bulk method the drying layer thickness should be much less for quick drying seeds than for slow drying seeds, such as peas (Pisum sativum L.) and beans (Phaseolus spp. L.). Chippindale (1934) reported that the imbibition of water in soaked and untreated seeds proceeds at about the same rate. Kidd and West (1918b) found the opposite to be 12 true. They stated that seeds soaked in a minimum amount of water and slowly dried imbibe water and develop more quickly than untreated seeds. Berrie and Drennan (1971) reported that imbibition in tomato seed was greater in soaked than in untreated seed.' Chippindale (1934) stated that since extreme desiccation after soaking does not remove the beneficial effect of the soaking, it is clear that the effect is independent of the absolute water content of the grains. Chippindale (1934) concluded that the accelera- tion of germination by presoaking is not associated directly with the water relations of the seed. The stage at which drying takes place after seed soaking, or even germination, appears to be the determining factor in plant survival according to Kydrev (1959). He also reported that drying exerts a favorable response on caryOpses which are at the stage of incipient germination, and plants grown from such caryopses develop most vigorously, accumulate the greatest amount of dry matter, and are most frost resistant. Berrie and Drennan (1971) showed that soaked tomato and oat (Aygna spp. L.) seeds could be returned to air dry condition provided that desiccation occurs before the onset of active cell division. Desiccation that occurred prior to cell division resulted in more rapid germination when the seeds were next wetted (Berrie and Drennan - 1971). The observation (Berrie and Drennan - 1971) that drying after cell division could be detrimental is supported by Kydrev (1959), who reported that Mercury wheat seeds were unable to 13 resume the dormant condition after formation of the coleoptile and emergence of the first leaf. Kydrev (1959) also showed that death of roots on germinating wheat seeds did not mean the seed (plant) would die, instead new roots often appeared shortly before the second soaking. Berrie and Drennan (1971) found that dehydration of tomato and oat seeds may be done more than one time, and the effects may be accumulative if prior imbibitions are of proper duration. The observation and conclusion that seed coverings do not return to their original state, thus leading to more rapid imbibition upon rewetting was reported by Berrie and Drennan (1971). Andersen (1938) indicated that two to three weeks of daily moistening and drying prior to germination gives a higher percent germination than untreated seeds under the same 20-30 C room temperature-germination environment. May, Milthorpe, and Milthorpe (1962) concluded that soaking seeds and then air drying produced plants which had increased drought tolerance. In dallisgrass (Paspalum dilatatum L.) the drying temperature did not appear to reduce seed viability, and in many cases seed dried at 60 C actually had stimulated germination to the point of being equal or better than seed dried at 37.8 C (Bennet and Marchbanks - 1969). Canode, Law, and Maguire (1970) found that Kentucky bluegrass seed which was dried in the swath, was able to withstand a temperature of 49 C without influencing germination. Ries, Seward, Schweizer, and Ayers (unpublished data) found that soaking seeds in organic solvents and then drying can increase the 14 growth and protein content of grains, under greenhouse and field conditions. Wheat seeds which were soaked in a large tank for 24 hours, then dried in.a kiln at 36-38 C to 12% moisture (approx. ten hours) had a positive yield increase only when the plants were subjected to stress, as reported by WOodruff (unpublished data). Bass (1953) reports that Kentucky bluegrass seed with 54% moisture was injured by 30 C drying for 24 hours, but 44% moisture seed withstood 60 C drying without loss in germination. Keller and Bleak (1968) and Bleak and Keller (1969) agree that although drying of seeds after being soaked produced more rapid germination rates than the control, there was still some loss in rate of emergence from the soaked seeds which were not dried. Several researchers have reported that in general, drying or certain drying techniques are ineffective in improving or retaining the advantage obtained by soaking seeds. Kidd and West (1918b) found that seeds which had been soaked and then dried rapidly germinate slower than untreated seeds. Waisel (1962) found that winter wheat which was soaked and dried or treated with calcium (Ca+) had no effect on true hardiness (frost, drought, and heat hardiness). Kreyger (1963) found that drying grains at temperatures greater than 65 C reduced seed viability. Finally, no ‘improvement in emergence rates of cotton, after soaking in water at several temperatures and then drying overnight at ambient temperature, were found by Thomas and Christiansen (1971). 15 Polyethylene Glycol (PEG) Soaking Treatment Heydecker, Higgins, and Guliver (1973) treated vegetable seeds with several soaking concentrations of PEG by placing the seeds in petri dishes on filter paper soaked in varying concentrations of the solutions. Following treatment, the seeds were washed, surface dried, and placed on water soaked filter paper to germinate. Heydecker et a1. (1973) stated that PEG cannot pass through the cell wall, produces an osmotic pressure, is chemically inert so it will not harm the seed during long term exposure, and its concentration is critical. Also, Heydecker et a1. (1973) reported that soaked onion (Allium cepa L.) seeds that were fully dried took three days to germinate in comparison to one day when not dried, yet even the drying treatment was significantly more rapid in germination than the control. The need for aeration during soaking was one problem which was also mentioned. Finally, the PEG treatments were found to increase both the rate and uniformity of germination, Heydecker et al. (1973). Field Results After Seed Soaking Treatments Adapting soaking treatments to a large scale for field establishment may present some potential problems due to volume. For example Kidd and West (1918b) reported that soaking seeds in large volumes of water may be harmful instead of stimulatory to seeds. Young et a1. (1974) noted difficulties with fermentation of the soaking liquid and seeds when hydroseeding. 16 Preplant soaking treatments of corn seeds showed a positive effect on the growth, development, and harvest of late planted corn according to Zubenko (1959). He also reported that the most effective treatments were those where single and double 24-hour soakings of seeds were followed by drying after each soaking to an air dry condition. Zubenko (1959) also found that corn seeds which sprouted during soaking had a lower germination than unsprouted seeds. Martyanova (1960). stated that boric acid soaking treatments appeared to be most effective during spring and summer droughts, after the appearance of shoots. New shoots of treated barley seeds appeared a day or two earlier than the control (Martyanova - 1960). Reduced germination at high osmotic pressures, simulated drought conditions, appears to be related to lower respiration and moisture absorption. Yet, improved germina- tion at high osmotic pressures was noted by Carry and Brown (1965) when wheat seeds were soaked prior to establishment. Following that line of thinking, Woodruff (1968) found no evidence of increased soil moisture usage by hardened wheat plants when compared to a control. Haight and Grabe (1972) noted up to a 40% increase in the rate of emergence from the soil of orchardgrass after a seed wetting and drying treatment. Bleak and Keller (1970) noted that with crested wheatgrass a soaked and a control did equally well in the greenhouse, while in the field the soaked seed emerged more rapidly than the untreated control. 17 .Abdel Hafeez and Hudson (1967) suggested that hardening, under some circumstances, may confer greater advantage under favorable conditions than under adverse growing conditions. In complete disagreement with Abdel Hafeez and Hudson (1967) are Chippindale (1934), Haight and Grabe (1972), and Bleak and Keller (1970) who feel that the advantage of preplant seed soaking was enhanced as conditions favoring emergence deteriorated. Germination Conditions In An Environmental Chamber Harrington (1923) found that bluegrass seed germinated most rapidly with alternations of temperature and also gave the most complete germination. Nelson (1927) also reported that alternating temperature is the most effective stimulant for the Egg spp. Toole and Toole (1955) reported that an alternating temperature between 15 C and 25 C was best for chewings fescue (Festuca rubra var. Commutata Gaud.), and that proper germination temperature is important in reducing variability of germination counts. Frazier (1960) stated that tall fescue and red fescue germinated under constant temperatures, but Penncross creeping bentgrass (Agrostis palustris L.) and Merion Kentucky bluegrass required alternating temperatures for maximum germination. Also, initial germination was accelerated by higher tempera- tures, but did not always produce the highest percentage of germination. Regarding chamber germination temperature, Haight and Grabe (1972) stated that an alternating temperature between 15 C and 25 C produced results for all treatments Is which were nearly equal in terms of their rates of seed germination, while at a 25 C constant germination tempera- ture the rates of germination were different for each treatment. In terms of light quantity during germination, Juhren, Hiesey, and Went (1953) found that Kentucky bluegrass did not benefit from high intensity, natural light. MATERIALS AND METHODS Determination of Optimum Soaking Time and Temperature Penncross creeping bentgrass (Agrostis palustris Huds.), Pennlawn red fescue (Festuca rubra L.), Merion Kentucky bluegrass (Poa pratensis L.), and Manhattan perennial ryegrass (Lolium perenne L.), were the four turfgrasses used in this study. Seed from the same four seed sources were used throughout the study. Twenty-five different soaking time- temperature treatments were established for each turfgrass species. Approximately 50 cc. of seed was placed in each soaking bottle. Soaking was accomplished in distilled water placed in 120 m1. glass bottles with screw-on plastic caps. Under these conditions, there appeared to be minimal air ‘ present during soaking, probably near anaerobic conditions since the water level in the bottles was kept filled to the lip, and the water was never changed. Soaking bottles were shaken each day to eliminate any air pockets, but were never aeriated while soaking. The temperature treatments were 5, 10, 15, 25 and 15-25 C, with the 15-25 C treatment alternating every 6 hours between 15 C and 25 C. Soaking times were 6, 12, 24, 48, and 168 hours. Soaking treatments took place in controlled environment chambers which were maintained in total darkness. 19 20 After the designated times, the appropriate bottles with the turfgrass seeds in them were removed from the soaking treatment chambers, at which time 25 seeds from each treatment bottle were placed in each of three petri dishes. This process of placing the seeds in the petri dish will be referred to as establishment throughout this paper. Petri dishes were new 10 cm plastic dishes which had #3 qualitative Whatman filter paper in each in order to maintain moist conditions. Once the seeds were placed in the petri dishes, the dishes, with covers, were placed in a large controlled environment chamber which was maintained at standard conditions for germination of these turfgrasses. These conditions were 8 hours of light at 25 C and 16 hours of darkness at 15 C. Both flourescent and incandescent lighting were present in the chamber which produced light energy ranging from 560-960 microeinsteins m"zsec-l within the chamber. Petri dishes were rotated weekly within the chamber, and were watered with distilled water as needed to maintain moist conditions. Germination counts were made on specific days, 5, 10, 15, 20, and 30 days after establishment in the germination chamber. The point at which the Shoot first emerged from the seed coat was defined as the point of germination for all four turfgrasses. Cgmparison of Oven vs. Air Drying Forty-five soaking temperature and time treatment combinations from the four turfgrasses studied in the above mentioned soaking study were dried in two different manners. 21 Soaking treatments took place exactly as described previously, then instead of placing the seeds in the petri dishes, the seeds from each bottle.were divided into two groups. One- half of the treated seeds, approximately 25 cc, were air dried on paper towels at an average temperature of 23.9 C for one week, while the other half was oven dried on paper toweling at 45 C for 18 hours. After drying treatments were complete, 25 seeds from each soaking and drying treatment combination were placed in - each of three new 10 cm plastic petri dishes with #3 Whatman qualitative filter paper. These dishes were then placed in the germination chamber with 8 hours of light at 25 C and 16 hours of darkness at 15 C for the 30 day observation period. Dishes were watered with distilled water as needed to maintain moist conditions within each dish. Germination readings were taken at times of 5, 10, 15, 20, and 30 days after establishment. Soaking in Polyethylene Glycol (Carbowax®>'6000') This study with Merion Kentucky bluegrass involving (a) two concentrations of polyethylene glycol (PEG), (b) several treatments soaked in distilled water, and (c) a control which was not soaked at all before established on the petri dishes, compared soaking treatments. Soaking time, soaking temperature, PEG concentration, aeration or no aeration, and the two previously mentioned drying techniques were the five treatment variables. 22 Soaking was done in one liter Erlenmeyer flasks with 200 cc of seed, at three times of 7, 12, or 21 days. Four temperature treatments were 10 C, 15 C, 23.9 C (room temperature), or alternating between 15 C and 25 C every six hours. The three concentrations were (1) 145 grams of CarbowaéE)'6000' in one liter of distilled water, (2) 290 grams of Carbowagat'GOOO' in one liter of distilled water, or (3) only pure distilled water. Aeration treatment was either aeriated or not aeriated. This was done with an impeller type pump which had a porous stone bubbler on the end of the air line within the soaking flasks. The aeration was intense enough to keep the solution and seed in constant motion. Drying treatments involved either air drying at 23.9 C for one week, oven drying at 45 C for 18 hours, or not drying at all before establishment. In all cases the seeds soaked in PEG were rinsed with distilled water to remove PEG on the seed surface before the 25 seeds were counted out into each of the three replicate petri dishes for the germination test. Controlled environment chambers maintained in total darkness were used for all soaking treatments, except for the room temperature treatments. Germination tests took place in another controlled environment chamber maintained at the standard germination conditions for these turfgrasses as stated previously. Also, distilled water was added to all petri dishes as needed to maintain moist conditions in 23 each dish. Germination observation times were the same: 5, 10, 15, 20 and 30 days after establishment. Field Study» Thirteen promising treatments from the three previous studies (soaking, drying, and PEG) were established, on September 20, 1974, in a split-plot design in the field on 152.4 cm X 152.4 cm plots, (5 X 5 ft. plots), with 4 replicates of each treatment. The plot area which had pH 6.6, phosphorus level of 70.62 Kg./ha. (63 lbs/A.) (high), and potassium level of 121.07 Kg./ha. (108 le/A.) (medium), was fertilized with 73.25 Kg./ha. (1.5 lbs/1,000 sq. ft.) of actual nitrogen (N) in the form of urea. This was applied with a gravity type spreader. Before seeding, the N was incorporated into the top 0.64 - 1.27 cm. (0.25 - 0.5 inch) of soil by a light raking. Another foliar application of urea at the rate of 24.41 Kg./ha. (0.5 1b/1,000 sq. ft.) was applied to the plots on November 9, 1974, to promote growth for the final field observations before snow cover. A Merion Kentucky bluegrass monostand was used in the field study. A seeding rate of 48.83 Kg./ha. (l lb/l,000 sq. ft.) was used for all plots. Seeding was done by hand within a perimeter seeding box, with the seed being mixed with sand for ease of distribution. Immediately after seeding, each plot was lightly raked to insure soil-seed contact in the upper .32 - .64 cm of soil, and then irrigated 24 lightly. Plots were neither rolled nor mulched. Alleyways were included to avoid walking on the actual plot area. Mulching would have made density measurement very difficult. Two replicates of the plot area were irrigated, with a rose-head nozzle, to maintain a moist soil surface, while the other half was only irrigated to maintain a moist soil surface for the first week after establishment. See Appendix Table 9 for rainfall and temperature data. Shoot density counts were taken in 10.16 X 10.16 cm steel grids on each plot, with three replicate random counts per plot, on both October 10 and October 15, 1974, 20 and 25 days, respectively, after establishment. Statistical Analysis The soaking, drying, and PEG studies were all completely randomized design within the growth chamber, while the field study was in a split-plot design. Means from all four studies were compared to the untreated control mean by using Dunnett's test at the 5% level. In all cases there were three replicates per each treatment, except in the field where there were four replicates. All percentages were transformed by are sine transformation before the analysis of variance and the Dunnett's test were performed. RESULTS AND DISCUSSION Results for all four turfgrasses are based on the germination percentages measured at 5, 10, 15, 20 and 30 days after establishment, with establishment being the time at which the seeds were placed in the petri dishes. Penncross creeping bentgrass Improvement in germination rate was noted only at the 5-day count (Table l and Appendix Table 1). After five days there was no positive effect of soaking on seed germination including the end point percent germination. There was also no effect from the five soaking temperatures. Soaking time was a significant factor. In general, increased soaking time (up to 168 hours as measured in this study) caused improved germination percentage. In all cases, in Table 1, where the soaking time was 48 hours or more, the 168 hour time gave higher percentage germination than the 48 hour time at the same soaking temperatures. Also worth noting is that the 24 C for 10 hours treatment was the only treatment significant at the 24 C temperature. Although inhibitory effects on seed germination were noted at the 5, 15, and 25 C levels where soaking time was 24 hours or less, the net temperature effect over the five measurement times was such that temperature was not a significant factor. 25 26 These soaking treatments may have an important effect on reducing damping-off problems, since the seeds could germinate fast enough to avoid being affected. No damping-off was noted in any phase of these studies. One of 14 treatments, 10 C soaking temperature for 48 hours and then air dried, gave significantly greater seed germination, and that was only at the 5-day observation date (Table 2 and Appendix Table 2). Detrimental effects of seed soaking followed by drying were noted primarily when the soaking temperature was greater than 15 C and soaked for 48 or 168 hours. Oven drying tended to be more inhibitory to early germination than air drying. The negative responses to drying seem to be related to the warmer soaking tempera- tures which may have induced respiration and cell division in the seed. Yet since the total germination percent was not effected, no reason for one positive response and several negative responses at the 5-day count has been hypothesized. Since no previous work with bentgrass was found, there is no means of direct species comparison to past results. This study shows that there is a definite advantage in terms of germination rate when Penncross creeping bentgrass seeds are soaked in distilled water for between 48 and 168 hours. Since the soaking temperature was not significant, this soaking technique could be useful and easily adapted to Penncross bentgrass establishment to insure more rapid emergence, and thus less potential for losses and poor stands due to erosion or weed infestation. The advantage obtained 27 by soaking Penncross seed was not retained after drying, whether by air or oven drying. Thus, the hydroseeding method appears to be the best means of utilizing the benefits from soaking bentgrass seed. These poSitive results with Penncross need to be compared with other bentgrass cultivars which have not been investigated. Pennlawn red fescue In this study there were two significant beneficial soaking treatments five days after establishment of the Pennlawn red fescue (Table 3 and Appendix Table 3). The best soaking treatments were for 48 hours or less. The temperature of the soaking treatments were not significant. Although, combining a soaking temperature of 25 C or 15-25 C with a soaking time of 168 hours resulted in permanent detri- mental effects on germination as well as the total end point germination. Possible explanation for the detrimental effects could be that the seeds began to respire at the higher temperature, lost some of the nutrients in the seed by leaching, and reduced the endosperm, through respiration and metabolism to a point where germination was no longer possible. As with bentgrass and most turfgrasses, little or no work has been done with soaking treatments on red fescue. Button (1959) and Behrendt (1960) have both worked with soaking red fescue with GA in the soaking solutions. Both found that the rate of emergence was hastened from a control. 28 Button and Noyes (1964) found that soaking red fescue with seaweed extract increased the rate of emergence. When Pennlawn was dried following various soaking treatments, there were no significant beneficial responses (Table 4 and Appendix Table 4). No significant differences were found between the air and oven drying techniques. The thirteen treatments which were negatively significant from the control were that way since the control reached 100% germination at the lO-day count, while the drying treatments reduced the germination percentage. Explanation for reduction in germination was not determined in this study. Soaking treatments that will increase seed germination of Pennlawn red fescue include 6 hours of soaking at 25 C, and 48 hours of soaking at 5 C. Most other soaking treat- ments with soaking times greater than 48 hours were permanently inhibitory to germination. In terms of drying, these results show that Pennlawn seed cannot be dried effectively after soaking without reducing the total end point percent germina- tion, thus making drying after soaking of Pennlawn seed impractical. Based on these findings, the previously listed beneficial soaking treatments would be best used in combination with hydroseeding. Manhattan perennial ryegrass In the Manhattan perennial ryegrass tests there were no soaking treatments nor soaking and drying combinations which were significantly better than an untreated control (Tables 5 and 6, and Appendix Tables 5 and 6). In the 29 soaking treatments, soaking temperatures of 25 and 15-25 C and a soaking time of 168 hours reduced total end point germination after 30 days. Six of the eight treatments in the drying study were significantly lower in percent germina- tion than the control. Over all five germination count times there was no significant difference between oven and air drying, but at the 5-day count there were significant differences between the air and oven techniques (Table 6 and Appendix Table 6). There are no past findings to relate to these results. Since perennial ryegrass germinates so rapidly without treatment, develOping a treatment to improve the rate of emergence may be of less importance. Manhattan perennial ryegrass may be losing necessary compounds for germination by leaching while soaking. This conclusion seems reasonable since most of the soaking and drying treatments have significantly lower germination percentages than the control. If drying was the cause of inhibition of germination then the soaking treatments should not have been affected as they were 0 Merion Kentucky bluegrass Soaking The length of soaking time was significant in the Merion Kentucky bluegrass seed soaking study (Table 7 and Appendix Table 7). The rate of germination generally increased with increasing soaking time. The only treatment combination significantly greater than the control was 30 soaking at 25 C for 168 hours, which was significant 10 days after establishment. Only three treatments were significantly lower than the control. These were only at the 20-day count and are therefore of little concern. Drying In the Merion drying study one treatment, 48 hours of soaking at 10 C and air dried, was’significantly greater than the control at the lS-day period only (Table 8 and Appendix Table 8). All other drying treatment combinations were far. from being positively significant compared to the untreated control. Most oven drying treatments caused significantly lower germination than the untreated control. Air drying gave significantly higher overall germination than oven drying. Polyethylene glycol (PEG) Soaking (PEG) treatments differed significantly from each other, but none positively from the untreated control (Appendix Table 10). Many treatments were inhibitory up to 20 days when compared to the control, after which no Signifi- cant differences existed from the control for all but 5 of the total 27 treatments. Although there were no positive effects in this study, the data seems to indicate: (a) oven drying ranked higher than air drying, (b) wet seed germinated more rapidly than air or oven dried seed, (c) PEG and distilled water performed equally, (d) 145 grams/liter and 290 grams/liter of water treatments responsed equally in 31 germination, and (e) aeriated treatments had a higher percent germination than non—aeriated soaking treatments. Field Study Results of the field study as determined by shoot density measurements made 20 and 25 days after establishment are reported in Table 9. Due toxa wet fall, the irrigated ya. non-irrigated comparisons were not significant at the 5% level as determined by an F-test. Visual differences between the 12 treatments and the untreated control were quite apparent in terms of green cover, but none of the treatments were significantly different in terms of density counts at the 5% level according to the F-test (Table 9). A possible explanation for why visual differences appear to exist, yet there is no significance after the statistical analysis of the density data is performed, is that the sampling error was larger than would be expected. Since the seed was established by hand there was difficulty in achieving a uniform stand on each plot. Thus greater than usual sampling error was encountered in taking the density counts. It should be noted that the best treatment from the growth chamber studies, 25 C soaking temperature with a soaking time of 168 hours and established wet, (see Table 7), was also one of the best ranking treatments in the field study (see Table 9). All other treatments from Tables 7 and 8 which were also in Table 9 responded in a consistent manner when compared between Tables. Therefore it appears that the 32 growth chamber is an effective means of evaluation and reducing seed soaking treatments before going to the field for confirming observations.i Had the fall not been so wet, it was hoped that any effects of soaking on drought tolerance and better emergence than untreated seed under suboptimal conditions would be determined. Thus further study is needed in the area of the effects of turfgrass seed soaking and drying where planted under suboptimal and stress conditions as discussed by Haight and Grabe (1972). Summary Discussion Rates of emergence can be increased in Penncross creeping bentgrass, Merion Kentucky bluegrass, and Pennlawn red fescue by soaking the seeds at a specific temperature and time combination in distilled water and sowing while wet. Penncross creeping bentgrass and Merion Kentucky bluegrass rates of emergence can also be improved by soaking in distilled water for a specific temperature and time combina- tion followed by air drying before sowing the seed. No soaking or drying treatment combination was found which would significantly increase the rate of emergence of Manhattan perennial ryegrass. From these studies it was concluded that no one treatment will be effective for all four of the turfgrasses studied, thus indicating that each species and cultivar will have its own unique stimulatory treatment. This conclusion is also supported by Young et al. (1974), Salim and Todd (1968), 33 Griswold (1936), Chippindale (1934), and Kidd and West (1918a and 1918b). Also, a possible explanation for some of the variability within treatments could be that single seeds within a source differ in reSponse to seed treatments, which ‘was suggested by the data of Keller and Bleak (1969) when working with preplant treatment of grass seed. Kidd and West (1919) stated that soaking in an excess amount of water was injurious at all temperatures, while in this study excess amounts of water were used for all soaking treatments with some positive results obtained. Also, this study showed that the rate of seed germination increases for some turfgrasses (Merion and Penncross) with increased soaking time which contradicts the results of Kidd and West (1919). Since it is not practical to report all daily germina— tion percentages, one is not able to determine from the Tables when each turfgrass initiated germination. Therefore, the following are ranges for initiation of germination of each turfgrass: Penncross creeping bentgrass 5—10 days, Manhattan perennial ryegrass 4-6 days, Pennlawn red fescue 5-8 days, and Merion Kentucky bluegrass 8-17 days. In regard to the drying treatments, oven and air dried, this research only reports the effect of oven drying for 18 hours at 45 C and air drying at 23.9 C for 1 week, not the general effect of all oven and all air drying treatments on seed which has been soaked. REFERENCES Abdel Hafeez, A.T., and J.P. Hudson. 1967. Effect of hardening radish seeds. Nature. 216:688. Andersen, A.M. 1935. The effect of soil in relation to nitrate solution on the germination of seed of Poa compressa. Amer. J. Bot. 22:906. . 1938. Moistening and drying as a pretreatment In germinating seeds of Poa compressa. Proc. Ass. Offic. Seed Anal. 30:246. . 1957. The effect of certain fungi and gibberellin on the germination of Merion Kentucky bluegrass seed. Proc. Ass. Offic. 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Jo 51:60-61. 34 35 Button, E.F., and C.F. Noyes. 1964. Effect of a seaweed extract upon emergence and survival of seedlings of creeping red fescue. Agron. J. 56:444-445. Canode, C.L., A.G. Law, and J.D. Maguire. 1970. Post- harvest drying rate and germination of Kentucky bluegrass seed. Crop Sci. 10:316-317. Chippindale, H.G; 1934. The effect of soaking in water on the "seeds" of some Gramineae. Ann. Appl. Biol. 21:225-232. Cole, D.F., and J.E. Wheeler. 1974. Effect of pregermination treatments on germination and growth of cottonseed at suboptimal temperatures. Crop Sci. 14:451-454. Curry, C.K., and R.H. Brown. 1965. Effects of various solutes at four osmotic concentrations on moisture absorption and germination of wheat. Agron. Abstr. p. 44. - Daniel, W.H. 1958. Why soak grass seed? Proc. Midwest Regional Turf Conf. pp. 30-31. ~ Daniel, W.H., and N.R. Goetze. 1957. Germination of turf cover species as influenced by periods of soaking in various fertilizer solutions. Agron. Abstr. p. 44. Dickens, R., and G. Moore. 1974. Effects of light, . temperature, KNO , and storage on germination of cogongrass. Agron. J. 66:187-188. Dunnett, C.W. 1955. A multiple comparison procedure for comparing several treatments with a control. J. Am. Stat. Assn. 50:1096-1121. Evenari, M. 1964. Hardening treatment of seeds as a means of increasing yields under conditions of inadequate moisture. Nature. 204:1010-1011. Frazier, S.K. 1960. Turfgrass seedling development under measured environment and management conditions. Master of Science Thesis, Purdue University. pp. 61. Frischknecht, N.C. 1959. Effects of presowing vernalization on survival and development of several grasses. J. ' Range Manag. 12:280-286. Genkel, P.A. 1964. Production experiments on the presowing hardening of plants against drought. (Russian), translated from Fiziologiya Rastenii. Vol. II, No. 3, pp. 538-543. 36 Griswold, S.M. 1936. Effect of alternate moistening and drying on germination of seeds of western range plants. Bot. Gaz. 98:243-269. Haight, J.C., and D.F. Grabe. 1972. Wetting and drying treatments to improve the performance of orchardgrass seed. Proc. Ass. Offic. Seed Anal. 62:135-148. Harrington, G.T.‘ 1923. Use of alternating temperatures in the germination of seeds. J. Agr. Res. 23:295-332. Heydecker, W., J. Higgins, and R. Gulliver. 1973. Bringing seeds to the brink of germination. The Grower (England). pp. 1134-1135. Juhren, M., W.M. Hiesey, and Went. 1953. Germination and early growth of grass in controlled conditions. Ecology. 34:288-300. Keller, W., and A.T. Bleak. 1968. Preplanting treatment to hasten germination and emergence of grass seed. J. Range Manag. 4:213-216. . 1969. Root and shoot growth following preplanting, treatment of grass seed. J. Range Manag. 22:43-46. Kidd, F., and C. West. 1918a. Physiological pre-determina- tion: The influence of the physiological condition of the seed upon the course of subsequent growth and upon the yield. I. The effects of soaking seeds in water. Ann. Appl. Biol. 5:1-10. . 1918b. Physiological pre-determination: The influence of the physiological condition of the seed upon the course of subsequent growth and upon the yield.. IV. Review of Literature. Chapter III. Ann. Appl. Biol. 5:220-251. . 1919. The influence of temperature on the soaking of’seeds. New Phytol. 18:35-39. Kreyger, J. 1963. General considerations concerning the drying of seeds. Int. Seed Testing Ass., Proc. 28:753-784. Kushnirenko, S.V. 1957. Physiological peculiarities of tomato and maize plants hardened to cold with alternating. temperature. U.S.S.R. Academy of Sciences, Moscow. pp. 233-241. Kydrev, T.G. 1959. The effect of presowing drought hardening of wheat caryopses on induced dormancy and germination. Bulgarian Academy of Sciences, Sofia, Bulgaria. pp. 179-183. 37 Linehan, P.A., and S.P. Mercer. 1936. Does pre-soaking accelerate laboratory germination in cocksfoot (Dactylis lomerata L.)? Int. Seed Testing Ass., Proc. Lyles, L., and C.D. Fanning. 1964. Effects of presoaking, moisture tension, and soil salinity on the emergence of grain sorghum. Agron. J. 56:518-520. Maguire, J.D., and K.M. Steen. 1971. Effects of potassium nitrate on germination and respiration of dormant and non-dormant Kentucky bluegrass (Poa pratensis L.) seed. Crop Sci. 11:48-50. Martyanova, K.L. 1960.__Resu1ts of field experiments with barley seed which had undergone a pre-sowing hardening to drought. (U.S.S.R.), translated from Fiziologiya Rastenii. vol. 7, No. 3. pp. 363-365. May, L.H., E.J. Milthorpe, and F.L. Milthorpe. 1962. Presowing hardening of plants to drought--an appraisal of the contributions by P.A. Genkel. Field Crop Abstr. 15:93-98. Nelson, A. 1927. The germination of Poa Spp. The Ann. Appl. Biol. 14(2):157-174. Parks, O.C., and P.R. Henderlong. 1967. Germination and seedling growth rate of ten common turfgrasses. Proc. West Virginia Acad. of Sci. Vol. 39. Porter, R.H. 1950. Relationship of hormones and inhibitors to seed germination. USGA J. 3(2):36. Ries, S.K., and C.J. Seward. Seed treatments of cereal grains to improve growth and protein content. II. Protein content and yield in greenhouse and field studies. (Unpublished data). Mich. State Univ. Salim, M.H. and G.W. Todd. 1968. Seed soaking as a pre-sowing, drought-hardening treatment in wheat and barley seedlings. Agron. J. 60:179-182. Thomas, R.I., and M.N. Christiansen. 1971. Seed hydration- chilling treatment effects on germination and subsequent growth and fruiting of cotton. Crop Sci. 11:454-456. Toole, E.H., and V.K. Toole. 1955. Variability in germina- tion as affected by physiological condition of seeds and germination microclimate. Proc. Ass. Offic. Seed Anal. p. 105. 38 Tyler, L.J., R.P. Murphy, and H.A. MacDonald. 1956. Effect of seed treatment on seedling stands and on hay yields of forage legumes and grasses. Phytopathology. 46:37-44. ‘ waisel, Yoav. 1962. Presowing treatments and their relation to growth and to drought, frost, and heat resistance. ' Physiologia Plantarum. 15:43-46. WOodruff, D.R. Evaluation of the pre-sowing drought hardening of wheat. (Unpublished data). Queensland Wheat Research Institute. . 1969. Studies on presowing drought hardening of wheat. Australian J. Agr. Res. 20:13-24. Young, J.A., B.L. Kay, and R.A. Evans. 1974. Plant growth regulators may enhance germination in hydroseeding. Plant Growth Reg. Bull. 2:19-20. Zubenko, V. Kh. 1959. The effect of preplanting hardening of seeds against drought on the grain harvest of corn in late plantings. (Krasnodar) Kuban Agricultural Institute. 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'MERION' AND 'KENBLUE' ABSTRACT Since previous research has shown that the growth of perennial ryegrass (Lolium perenne L.) to be increased by foliar applications of simazine and atrazine, and also since foliar applications of urea on wheat (Triticum aestivum L.) increased yield and protein content, this study was undertaken to determine what effect such foliar applications would have on the rate of germination of turfgrass seeds formed under the influence of such treatments. Kentucky bluegrasses (Poa pratensis L.) 'Merion' and 'Kenblue' were treated with foliar applications of simazine and urea with both one and three application treatments. The first application to the inflorescence was at anthesis, with the second and third applications being seven and fourteen days after anthesis. Identical treatments were made with and without using a wetting agent. The results indicate that none of the treatment combinations with urea, simazine, and a wetting agent were significantly better than the control in terms of percent seed germination at 5, 10, 15, 20, and 30 days after establishment, according to Dunnett's test for two-sided comparisons at the 5% level. 49 INTRODUCTION Kentucky bluegrass (Poa pratensis L.) is notorious for a relatively slow germination rate compared to most other cool season turfgrasses. Attempts have been made at finding a method of reducing the time of emergence of Kentucky blue- grasses Andersen (1957), Bass (1953), Behrendt (1960), Daniel and Goetze (1957), Daniel (1958), Harrington (1923), Maguire and Steen (1971). Germination requirements of a given species or cultivar vary depending on conditions during development, on maturity, and on age of seed, according to ToOle and Toole (1955). Sub-herbicidal rates of atrazine and simazine have been reported by Kay (1969) to double yields, and also to increase protein and nitrate nitrogen in wheatgrass seed (Agropyron intermedium (Host) Beauv.), as well as increase growth and protein content of perennial ryegrass (Lolium perenne L.). An application rate of 1.121 Kg./ha. was reported most effective. Finey, Meyer, Smith, and Fryer (1957) found that foliar spray applications of urea consisting of one to several applications before and during anthesis caused higher yields and increased protein content in the seed of Pawnee wheat (Triticum aestivum L.). The effect of multiple sprayings was not additive, Finney et a1. (1957). 50 51 The objectives of this study were to determine (1) if a foliar spray application of urea during seed development at one or three applications, with and without wetting agents, 'would promote development of a more vigorous seed in terms of rate of shoot emergence, and (2) if sub-herbicidal rates of simazine applied during seed development would promote development of a seed which had a more rapid rate of shoot emergence. Improving emergence rates of Kentucky bluegrass could possibly eliminate the need for mixing rapid germinating species with Kentucky bluegrass when seeding to prevent soil erosion, reduce weed infestation during establishment, and even replace the use of sod in some instances. MATERIALS AND METHODS Field Site Description Mature stands of Kentucky bluegrass (Poa pratensis L.) 'Merion' and 'Kenblue' grown on a Houghton muck soil were used for this investigation. The uniformity and density of stand and seedhead formation were excellent for Kenblue and good for Merion. These two blocks were set up in a completely randomized block design comprising four replicates of each treatment. Plot size was 1.82 X 1.52 m. for Kenblue and 1.98 X 2.13 m. for Merion. Phosphorus and potassium levels were medium for turfgrass culture, with the soil having a pH 6.9. Nitrogen was not applied during the study, other than in the treatments described, or in the year previous to the study. No irrigation was used in the past years nor during the investigation. Finally, the plot area had been mowed once per year for the past four years. Foliar Spray Applications The eight inflorescence treatments included: (a) one application of 56.05 kg. urea/ha. (150.13 g. of urea/liter of water) applied with wetting agent at anthesis, (b) one application of 33.63 kg./ha. of urea (90.08 g. of urea/liter of water) applied with wetting agent at anthesis, (c) one application of 56.05 kg./ha. of urea applied without wetting 52 53 agent at anthesis, (d) one application of 33.63 kg./ha. of urea applied without wetting agent at anthesis (e) three applications of 56.05 kg./ha. of urea applied with wetting agent at anthesis and at Seven and fourteen days after anthesis, (f) three applications of 33.63 kg./ha. of urea applied with wetting agent at anthesis and at seven and fourteen days after anthesis, (g) one application of 0.28 kg. a.i. of simazine per hectare applied at anthesis, and (h) one application of 0.07 kg. a.i. of simazine per hectare applied at anthesis. Wetting agent, Adjuvan-‘I®, was applied at a concentration of 25.43 liters/ha. The simazine was 80% A.I. Four replications of an untreated control were maintained for each of the two Kentucky bluegrasses. Spray applications were applied with a 3.78 liter capacity, hand pump, pressure sprayer which had a boom with two nozzles giving a 2.0 meter spray width. Calibration of the sprayer was performed prior to each application. Spray volume was 570 liters of water/ha. (60.5 ga1./A.). Conditions during spraying were partly cloudy to clear, 1-7 mph. winds, and from 15.6 - 28.3 C. Precipitation of 0.5 cm. occurred four hours after the first application, June 13, 1974, with no precipitation measured for 48 hours after the second and third applications, June 20 and June 27, 1974. Harvest and Drying, Harvest took place on July 10, 1974 when the seed heads were golden brown and the seed was just beginning to fall. The peduncles with the spikes were harvested from each 54 plot and placed in paper bags which were placed in the greenhouse to dry for two months. ‘The seed was then removed by rubbing the spikelets between the fingers, and afterwards was placed in sealed envelopes. Germination Test Germination evaluations were conducted in new 10 cm. plastic petri dishes containing #3 qualitative Whatman filter paper. Each dish received water as needed to maintain moist conditions at all times. Eight petri dishes with 25 seeds per dish were made for each treatment, comprised of fdur replicates with two subsamples per replicate. The optimal germination conditions used were 18 hours of darkness at 15 C and 6 hours of light at 25 C (Harrington - 1923), and Nelson (1927), which were maintained in a controlled environment chamber. Germination was defined as the point at which the shoot first emerged from the seed coat. Observations were made at 5, 10, 15, 20, and 30 days after placement in the petri dishes. Statistical Analysis A completely randomized analysis of variance was made on each of the two grasses, with a two-sided Dunnett's test performed at the 5% level of significance on the means. Each mean was made up of eight germination evaluation percentages. Also, all percentages were transformed by arc sine transformation prior to the analysis of variance and Dunnett's test. RESULTS AND DISCUSSION Merion Kentucky bluegrass None of the inflorescence treatments were significantly different from the untreated control at the 5% level using Dunnett's test (Dunnett 1955) for two-sided comparisons (see Table 10). Yet, several of the treatments did produce results which ranked higher than the untreated control at the 10 and 15 day observation times. There is no literature to refer to on the effect of foliar applications of simazine and urea on the germination rate of Merion Kentucky bluegrass. Kenblue Kentucky bluegrass Some of the inflorescence treatments on Kenblue Kentucky bluegrass were significantly different at the 5% level, yet none of the treatment means were significantly different from the untreated control at the 5% level using Dunnett's test for two sided comparisons (see Table 11). As with Merion, several of the treatments did produce results which ranked higher than the control, in this case at 10, 15, 20, and 30 day observation times. Again, since Similar work has never been published, there is no means of comparison with this study. In conclusion, the effects of foliar applications of urea and simazine on Merion and Kenblue Kentucky bluegrass do 55 56 not seem to have a place in terms of improving the rate of emergence of these turfgrasses under optimal conditions in a growth chamber. This study does confirm that the effect of multiple sprayings was not additive, as previously stated by Finney et a1. (1957). The effects of multiple applications, varied concentrations of both simazine and urea, and a wetting agent did not prove to be significantly effective in terms of improving the germination rates. Further studies are needed to determine yield, seed size, protein content, field germination response, and optimum treatment, if any, after similar treatments as described in this study. 57 TABLE 10. Effects of foliar applications of urea and simazine on the germination rate of Merion Kentucky bluegrass. Treatments (as described in materials and methods % Germination: 10,15,20,and 30 days after establishment (means of 8 replications) 10 day . 15 day 20 day 30 day Control 45.00 72.00 81.50 84.00 C 45.50 65.00 75.00 78.00 A. 40.50 55.00 69.50 72.50 E 42.00 61.00 69.50 71.50 D 26.00 51.00 66.50 74.38 B 46.00 74.50 79.50 83.50 F 49.50 69.00 75.00 79.50 H 25.00 66.50 77.00 78.50 G 58.00 73.00 80.00 81.00 NOne of the above means are significantly different from the control at the 5% level using Dunnett's test for two-sided comparisons. 58 TABLE 11. Effects of foliar applications of urea and simazine on the germination rate of Kenblue Kentucky bluegrass. Treatments % Germination: 10,15,20,and 30 days (as described in materials and methods) after establishment (means of 8 replications) 10 day 15 day 20 dav ‘30 day Control 31.75 55.50 64.50 67.00 c 29.00 63.50 73.50 78.50 A 44.50 65.60 72.00 77.00 E 47.00 65.50 74.50 79.00 D 39.00 60.50 68.38 74.88 B 23.00 42.00 54.00 63.00 F 24.50 48.50 66.50 71.00 H 28.50 48.50 60.50 64.00 G 26.50 55.00 63.00 68.00 None of the above means are significantly different from the control at the 5% level using Dunnett's test for two-sided comparisons. REFERENCES Andersen, A.M. 1957. The effect of certain fungi and gibberellin on the germination of Merion Kentucky bluegrass seed. Proc. Ass. Offic. Seed Anal. 47:145. Bass, L.N. 1953. Relationship of temperature, time, and moisture content to viability of seeds of Kentucky bluegrass. Iowa Acad. Sci. 60:86-88. Behrendt, S. 1962. Possibilities of acceleration of the germination of perennial grass species by gibberellic acid. Eigenschaften Wirkungen Gibberelline, Symp. Giessen, Germany, 1960. pp. 195-199. Daniel, W.H., and N.R. Goetze. 1957. Germination of turf cover species as influenced by periods of soaking in various fertilizer solutions. Agron. Abstr. p. 44. Daniel, W.H. 1958. Why soak grass seed? Proc. Midwest Regional Turf Conf. pp. 30-31. Dunnett, C.W. 1955. A multiple comparison procedure for comparing several treatments with a control. J. Am. Stat. Assn. 50:1096-1121. Finney, K.F., J.W. Meyer, F.W. Smith, and H.C. Fryer. 1957. Effect of foliar spraying of Pawnee wheat with urea solutions on yield, protein content, and protein quality. Agron. J. 49:341-347. Harrington, G.T. 1923. Use of alternating temperatures in the germination of seeds. Agr. Res. 23:295-332. Kay, Burgess L. 1969. The increase in protein content and yield of simazine and atrazine treated range forage. (Unpublished data). California. Maguire, J.D., and K.M. Steen. 1971. Effects of potassium nitrate on germination and respiration of dormant and non-dormant Kentucky bluegrass (Poa pratensis L.) seed. 59 f CONCLUSIONS The following conclusions are drawn based on the results of these investigations: 1. V Germination of each turfgrass studied responded differently to seed soaking and drying treatments, indicating that no one treatment combination would be effective for all species or cultivars. The longer the seed soaking time (up to 168 hrs.) for Penncross creeping bentgrass and Merion Kentucky bluegrass, the more rapid were the rates of shoot emergence. Seed soaking temperature was significant only with Manhattan perennial ryegrass. Seed soaking times of less than 48 hours were found to be best for Pennlawn red fescue. No seed soaking and drying treatments were found which could significantly (5% level) improve the rate of shoot emergence of Manhattan perennial ryegrass. Treatments were found which did significantly (5% level) increase the rate of emergence of Penncross creeping bentgrass, Pennlawn red fescue, and Merion Kentucky bluegrass. 60 10. 11. 61 The optimal soaking treatments found for Penncross creeping bentgrass were 5 C for 168 hours, 15 C for 168 hours, and 15-25 C for 168 hours, with 10 C for 48 hours then air dried being the best soaking-drying combination. For Pennlawn red fescue, the two highest ranking soaking treatments were 25 C for 6 hours and 5 C for 48 hours, while no soaking-drying treatment had positive results. Merion Kentucky bluegrass had an optimal soaking treatment of 25 C for 168 hours. The soaking- drying treatment of 10 C for 48 hours and then air dried showed the most promise. In the field, 15 C for 6 hours and 15-25 C fOr 12 hOurs seeded wet resulted in the most dense stands. Polyethylene glycol soaking treatments were not effective at any temperature, soaking time, or concentration. The soaking effect on germination was inhibitory for many treatments. Also, although there were no positively significant treatments compared to the control, the treatments which were aeriated resulted in a greater percent germination than the non—aeriated soaking treatments. Soaking in a large quantity of water is not. necessarily detrimental to the seeds, as was previously indicated by Kidd and West (1919). 12. 13. 62 Air drying at 23.9 C after soaking was significantly better (5% level) than oven drying at 45 C after soaking, with Penncross creeping bentgrass and Merion Kentucky bluegrass. With Manhattan perennial ryegrass and Pennlawn red fescue, oven drying after soaking was more detrimental to germination than air drying. Liquid applications of simazine and urea to the inflorescence, with and without a wetting agent, during and after anthesis, did not have any significant (5% level) effect upon the subsequent rate of shoot emergence of Merion and Kenblue Kentucky bluegrasses. APPENDIX 63 APPENDIX TABLE 1. Penncross creeping bentgrass soaking treatment means for seed germination. Percent Seed Germination at 5 dates after sowing the seed Treatment (means of 3 replicates) Saning Time Soaking‘Temp. TDays) (hrs.) (°C) 5 10 15 20 30 6 5 8.0 86.7 90.7 96.0 96.0 6 10 14.7 73.3 89.3 89.3 89.3 6 15 1.3 66.7 93.3 96.0 96.0 6 25 0 69.3 92.0 93.3 97.3 6 15-25 9.3 68.0 82.7 94.7 94.7 12 5 1.3 90.7 100. 100. 100. 12 10 8.0 76.0 93.3 96.07 97.3 12 15 0 61.3 98.7 98.7 98.7 12 25 6.7 80.0 89.3 93.3 93.3 12 15-25 6.7 64.0 92.0 94.7 98.7 24 5 10.7 66.7 89.3 94.7 98.7 24 10 58.7 93.3 98.7 98.7 98.7 24 15 16. 86.7 89.3 92.0 96.0 24 25 13.3 94.7 98.7 98.7 98.7 24 15-25 38.7 94.7 98.7 98.7 98.7 48 5 57.3 94.7 96.0 96.0 96.0 48 10 66.7 96.0 96.0 97.3 97.3 48 15 73.3 94.7 96.0 96.0 96.0 48 25 44.0 89.3 93.3 94.7 94.7 48 15-25 58.7 96.0 96.0 96.0 96.0 168 5 81.3 94.7 94.7 94.7 94.7 168 10 69.3 85.3 86.7 86.7 86.7 168 15 81.3 93.3 94.7 96.0 96.0 168 25 77.3 98.7 98.7 98.7 98.7 168 15-25 81.3 98.7 98.7 98.7 98.7 Untreated Control 15.0 89.3 94.7 97.3 98.7 64 APPENDIX TABLE 2. Penncross creeping bentgrass soaking- drying treatment means for seed germination. Percent Seed Germination at __ Tpeatment 5 dates after sowing the seed Soaking Soaking (means of 3 replicates) Time Temp. Drying Ifiéys) (hrs.) (°C) Treatment 5 10 15 20 30 12 5 air 49.3 98.7 98.7 98.7 98.7 12 5 oven 54.7 100 100 100 100 24 10 air 68.0 100 100 100 100 24 10 oven 41.3 100 100 100 100 24 25 air 48.0 98.7 98.7 98.7 98.7 24 25 oven 58.7 98.7 100 100 100 24 15-25 air 42.7 96.0 97.3 97.3 97.3 24 15-25 oven 42.7 97.3 97.3 98.7 98.7 48 5 air 61.3 100 100 100 100 48 5 oven 62.7 100 100 100 100 48 10 air 89.3 100 100 100 100 48 10 oven 66.7 98.7 98.7 98.7 98.7 48 15 air 65.3 94.7 96.0 96.0 96.0 48 15 oven 46.7 100 100 100 100 48 25 air 56.0 90.7 92.0 93.3 93.3 48 25 oven 44.0 96.0 96.0 96.0 96.0 48 15-25 air 74.7 97.3 98.7 100 100 48 15-25 oven 32.0 94.7 96.0 96.0 96.0 168 5 air 88.0 97.3 98.7 98.7 98.7 168 5 oven 45.3 93.3 100 100 100 168 10 air 76.0 92.0 92.0 92.0 92.0 168 10 oven 50.7 97.3 97.3 97.3 98.7 168 15 air 58.7 98.7 98.7 98.7 98.7 168 15 oven 44.0 98.7 98.7 98.7 98.7 168 25 air 34.7 88.0 90.7 92.0 92.0 168 25 oven 26.7 82.7 88.0 89.3 89.3 168 15-25 air 48.0 97.3 97.3 97.3 97.3 168 15-25 oven 29.3 97.3 98.7 98.7 98.7 Untreated Control 64.0 96.0 96.0 96.0 96.0 65 APPENDIX TABLE 3. Pennlawn red fescue soaking treatment means for seed germination. Percent Seed Germination at 5 dates after sowing the seed Treatment (means of 3 replicates) Saning Time Soaking Temp. (Days) (hrs.) (°C), 5 10 15 20 30 6 5 0 93.3 94.7 94.7 94.7 6 10 O 89.3 90.7 92.0 92.0 6 15 1.3 88.0 97.3 98.7 98.7 6 25 12.0 90.7 94.7 94.7 94.7 6 15-25 0 89.3 97.3 97.3 97.3 12 5 0 89.3 96.0 97.3 98.6 12 10 1.3 92.0 94.7 94.7 94.7 12 15 O 88.0 94.7 94.7 94.7 12 25 0 87.3 97.3 97.3 97.3 12 15-25 0 92.0 93.3 94.7 94.7 24 5 1.3 86.7 97.3 98.7 98.7 24 10 6.7 90.7 96.0 96.0 97.3 24 15 0 92.0 97.3 98.7 98.7 24 25 2.7 82.7 97.3 97.3 97.3 24 15-25 0 86.7 93.3 94.7 96.0 48 5 8.0 92.0 93.3 94.7 96.0 48 10 5.3 92.0 96.0 96.0 96.0 48 15 2.7 92.0 97.3 97.3 97.3 48 25 2.7 86.7 94.7 97.3 97.3 48 15-25 1.3 88.0 94.7 94.7 97.3 168 5 0 94.7 98.7 98.7 98.7 168 10 2.7 92.0 98.7 98.7 98.7 168 15 4.0 77.3 89.3 92.0 92.0 168 25 1.3 66.7 78.7 81.3 82.7 168 15-25 0 65.3 76.0 81.3 81.3 Untreated Control 0 96.0 98.7 98.7 98.7 66 APPENDIX TABLE 4. Pennlawn red fescue soaking-drying treatment means for seed germination. Percent Seed Germination at _g_ Treatment 5 dates after sowing the seed Soaking Soaking~ (means of 3 replicates) Time Temp. Drying (Déys) (hrs.) (°C) Treatment 5 10 15 20 30 6 25 air 1.3 93.3 97.3 100 100 6 25 oven 0 96.0 97.3 98.7 98.7 24 10 air 14.7 97.3 98.7 98.7 98.7 24 10 oven 0 98.7 98.7 98.7 98.7 48 5 air 10.7 97.3 97.3 98.7 98.7 48 5 oven 0 86.7 89.3 90.7 90.7 48 10 air 37.3 94.7 98.7 98.7 98.7 48 10 oven 29.3 97.3 98.7 98.7 98.7 48 25 air 4.0 92.0 93.3 93.3 93.3 48 25 oven 0 89.3 93.3 93.3 93.3 168 10 air 24.0 89.3 92.0 92.0 93.3 168 10 oven 25.3 90.7 92.0 92.0 92.0 168 15 air 1.3 82.7 85.3 86.7 88.0 168 15 oven 0 89.3 93.3 94.7 94.7 Untreated Control 13.3 100 100 100 100 67 APPENDIX TABLE 5. Manhattan perennial ryegrass soaking treatment means for seed germination. Percent Seed Germination at 5 dates after sowing the seed Treatment" (means of 3 replicates) SoERing Time Saning Temp. TDayS) (hrs.) (°C) 5 10 15 20 30 6 5 5.3 89.3 93.3 94.7 94.7 6 10 16.0 86.7 94.7 94.7 94.7 6 15 13.3 88.0 93.3 93.3 94.7 6 25 12.0 76.0 84.0 85.3 85.3 6 15-25 10.7 88.0 94.7 94.7 94.7 12 5 22.7 80.0 86.7 89.3 89.3 12 10 52.0 90.7 90.7 92.0 92.0 12 15 33.3 90.7 90.7 90.7 90.7 12 25 24.0 88.0 93.3 94.7 94.7 12 15-25 37.3 98.7 98.7 98.7 98.7 24 5 41.3 96.0 97.3 98.7 98.7 24 10 30.7 92.0 96.0 96.0 96.0 24 15 25.3 94.7 97.3 97.3 97.3 24 25 9.3 93.3 98.7 98.7 98.7 24 15-25 20.0 93.3 96.0 96.0 96.0 48 5 22.7 92.0 96.0 96.0 96.0 48 10 28.0 88.0 93.3 93.3 94.7 48 15 34.7 90.7 94.7 94.7 94.7 48 25 13.3 76.0 89.3 92.0 93.3 48 15-25 22.7 93.3 100. 100. 100. 168 5 24.0 96.0 96.0 100. 100. 168 10 21.3 96.0 97.3 97.3 100. 168 15 21.3 78.7 93.3 93.3 93.3 168 25 O 41.3 76.0 82.7 82.7 168 15-25 2.7 66.7 89.3 92.0 93.3 Untreated Control 28.0 94.7 96.0 97.3 97.3 68 APPENDIX TABLE 6. Manhattan perennial ryegrass soaking- drying treatment means for seed germination. Percent Seed Germination at Treatment 5 dates after sowing the seed Soaking Soaking (means of 3 replicates) Time Temp. Drying (Days) (hrs.) (°C) Treatment 5 10 15 20 30 12 10 air 29.3 97.3 97.3 97.3 97.3 12 10 oven 13.3 92.0 92.0 92.0 92.0 12 15 air 29.3 93.3 94.7 94.7 94.7 12 15 oven 10.7 90.7 92.0 92.0 92.0 12 15-25 air 26.7 93.3 93.3 93.3 94.7 12 15-25 oven 18.7 98.7 100 100 100 24 5 air 48.0 96.0 97.3 98.7 100 24 5 oven 8.0 96.0 97.3 97.3 98.7 24 10 air 34.7 97.3 98.7 98.7 98.7 24 10 oven 16.0 93.3 94.7 94.7 94.7 48 15 air 28.0 92.0 94.7 94.7 94.7 48 15 oven 10.7 98.7 98.7 98.7 98.7 48 15-25 air 29.3 96.0 97.3 97.3 97.3 48 15-25 oven 8.0 92.0 98.7 98.7 98.7 168 10 air 60.0 94.7 94.7 94.7 94.7 168 10 oven 58.7 94.7 94.7 94.7 94.7 Untreated Control 77.3 97.3 97.3 97.3 97.3 69 APPENDIX TABLE 7. Merion Kentucky bluegrass soaking treatment means for seed germination. Percent Seed Germination at 5 dates after sowing the seed Treatment . (means of 3 replicates) Saning Time Soaking Temp. (Days) (hrs.) (°C) 5 10 15 20 30 6 5 0 1.3 6.7 29.3 58.7 6 10 0 0 5.3 36.0 66.7 6 15 0 5.3 33.3 53.3 73.3 6 25 0 0 16.0 50.7 76.0 6 15-25 0 0 17.3 40.0 77.3 12 5 O 1.3 10.7 41.3 69.3 12 10 0 5.3 17.3 36.0 68.0 12 15 0 1.3 14.7 45.3 70.7 12 25 0 0 6.7 25.3 50.7 12 15-25 0 4.0 24.0 52.0 73.3 24 5 O 0 5.3 44.0 82.7 24 10 0 5.3 16.0 33.3 78.7 24 15 0 8.0 30.7 64.0 85.3 24 25 0 1.3 16.0 50.7 78.7 24 15-25 0 6.7 26.7 68.0 85.3 48 5 0 4.0 29.3 60.0 65.3 48 10 0 2.7 21.3 57.3 80.0 48 15 0 1.3 17.3 52.0 76.0 48 25 0 4.0 17.3 36.0 61.3 48 15-25 0 2.7 20.0 46.7 76.0 168 5 O 2.7 33.3 68.0 84.0 168 10 0 1.3 18.7 42.7 84.0 168 15 0 9.3 24.0 42.7 85.3 168 25 0 10.7 36.0 61.3 81.3 168 15-25 0 5.3 29.3 54.7 85.3 Untreated Control 0 0 14.7 57.3 73.3 70 APPENDIX TABLE 8. Merion Kentucky bluegrass soaking-drying treatment means for seed germination. Percent Seed Germination at ‘_ Treatment 4 dates after sowing the seed SoiEing Soaking- (means of 3 replicates) Time Temp. Drying —TDays) (hrs.) (°C) Treatment 10 15 20 30 6 15 air . 1.3 34.7 57.3 80.0 6 15 oven 2.7 26.7 62.7 80.0 12 10 air 2.7 22.7 57.3 74.7 12 10 oven 0 18.7 46.7 72.0 12 15-25 air 4.0 20.0 66.7 86.7 12 15-25 oven 1.3 18.7 61.3 89.3 24 10 air 2.7 44.0 78.7 86.7 24 10 oven 2.7 22.7 66.7 80.0 24 15 air 5.3 45.3 62.7 74.7 24 15 oven 2.7 18.7 50.7 76.0 24 15-25 air 2.7 37.3 73.3 92.0 24 15-25 oven 1.3 9.3 36.0 73.3 48 5 air 6.7 38.7 66.7 82.7 48 5 oven 5.3 36.0 66.7 78.7 48 10 air 24.0 74.7 86.7 93.3 48 10 oven 8.0 46.7 70.7 85.3 48 15 air 0 36.0 66.7 80.0 48 15 oven 8.0 32.0 66.7 82.7 48 25 air 2.7 37.3 65.3 86.7 48 25 oven 4.0 20.0 52.0 68.0 48 15-25 air 2.7 30.7 57.3 84.0 48 15-25 oven 1.3 9.3 32.0 69.3 168 5 air 9.3 50.7 78.7 89.3 168 5 oven 1.3 32.0 54.7 85.3 168 10 air 9.3 44.0 70.7 88.0 168 10 oven 2 21.3 52.0 84.0 168 15 air 1 24.0 46.7 72.0 168 15 oven 0 13.3 44.0 82.7 168 25 air 2 22.7 58.7 81.3 168 25 oven 1 5.3 26.7 54.7 168 15-25 air 1 10.7 49.3 73.3 168 15-25 oven 2 9.3 30.7 70.7 Untreated Control 5 46.7 72.0 85.3 71 APPENDIX TABLE 9. Temperature and rainfall data from September 20 to October 15, 1974, in East Lansing, Michigan, as recorded by the U.S. Dept. of Commerce Climatological Data. Date Rainfall (cm.) Temperature (°C) minimum maximum average September 20 .05 5.6 16.1 11.1 21 .08 5.0 17.8 11.7 22 T -5.6 11.1 5.6 23 O —2.2 15.0 6.7 24 T 7.2 17.8 12.8 25 T 2.2 19.4 11.1 26 O 3.3 27.8 15.6 27 1.29 11.7 25.6 18.9 28 .58 16.7 23.9 20.6 29 1.93 5.0 18.3 11.7 30 2.13 5.0 9.4 7.2 October 1 .10 .6 10.0 5.6 2 .20 -3.9 1.4 1.1 3 O -5.6 11.7 3.3 4 T 3.2 20.6 13.9 5 T 11.1 23.3 17.2 6 .41 6.1 22.8 14.4 8 O -2.2 15.6 6.7 9 O 1.7 15.0 8.3 10 O 1.7 21.7 11.7 11 O 3.3 22.8 13.3 12 T 4.4 17.2 11.1 13 .58 -2.2 14.4 6.1 14 1.09 8.3 17.2 12.8 15 O 0 12.2 5.6 72 o.~n s.mm s.om4 e.~ a oouno nuoxoouounos\omo .ooomxomme p.4e m.asa s.o~s s.~ 4 oouno uuo\oououno<\omo .oOsH\HN\OH m.oo n.oos S.OH. m.o a oouno so>o\oououno<\omo .oOeH\mH\usouns< M.Ho o.oma ~.o~4 o.o oouno so>o\oouounos\omo .omeo\um\oo s.oos s.ems o.mHs o a oouno oo>o\oouounomxnouos ooHHHunHO\e\O~-mH M.HO o.so m.ema o.~H oouno oo>o\oouounoe uo2\wmn .omsH\o~\OH m.es m.em s.oms m.m oouno uuo\ooununo<\nouoz ooHHHunuo\e\mmumo o.om n.4ms m.o~4 m.O mouse uuo\oououuo< uo2\omn .oOeH\H~\OH m.me m.mos m.eHs o s oouno nuo\oouounoe\oma .omeH\H~\oH m.se m.oss o.e~4 m.m 4 oouno nuo\oouounos\oma .oOsH\~H\usouos< m.aO a.~o m.em4 o.OH Dos ooooomxoououno<\oma .oOsH\~H\us8uos< s.~o m.mo m.so o.- Honusoo oouoonuso s.oo m.oo o.os m.eH no: ooooomxoououno<\omm .oOsH\H~\oH e.os o.oo M.Ho, M.HN nos ooooom\oouounos uo2\omn .oOeH\HN\OH m.me m.mm e.sms M.HN nos ooooom\oouounos\uouo3 ooHHuunuoxexmmumH o.OO o.mo o.so a.eH no; ooooom\oouounos\omn .oOsH\H~\OH o.~o m.OO m.mo n.4H uoz ooooom\oououno<\omo .ooo~\H~\OH o.om m.sm o.~m4 s.~ . nos ooooom\oouounos\omn .Ooo~\~H\usoHnsm o.mm M.Ho o.~ma o.s . nos ooooom\oouounos uo2\omo .Ooo~\H~\mH o.~o moms o.mo e.om no; ooooom\oouounos\omo .oOsH\n\O~uOH om om OH oH souuuosoo ooom\oouuounom\souuoHom mmmo HmsumOHHmwu O HO mCmsEv .mossm sou OCHBOO usumm msumo v um CoHumCHEuso ossm quOHso \Amhmov sEHB wmommaony mass xmom quEumsuB ossm .mCmsE CoHumCHEusm quEumsuu OCmeOO ossm HOOmHm sCsHOousmHoo .OH mnmde NHQZNmmd 73 m.uusCCso .mCOmHHmoEoo osoHOIOSu H0O umsu OCHmC Hs>sH OO sou um HouuCos sou Cmou COHumCHEHsm quouso ussoH aHqusHmHCmHm4 H.NO4 m.mm4 H.4H4. m.H 4 ooHno soooxoouounoa\omo .oom~\~H\uoOHns< M.HO o.om4 m.mm4 m.O oouno so>o\oououuos\omm .oOoH\s\O~rOH o.oo4 m.H~4 H.O 4 o 4 OOHno so>oxoouounoa uo2\omo .Ooam\Hm\mH m.oo4 o.~m4 a.oH4 o 4 ooHno nHoxoouoHnom noz\omm .Oom~\H~\OH o.eO4 o.~m4 o.~H4 e.~ 4 OOHno nuoxoouounoaxomn .oOoH\n\OmuOH m.om n.ms H.4O o.O ooHno so>o\oououuos\omn .oomm\H~\OH o.om o.oo s.mm4 n.4H oouno so>o\oouounom\omn .oOsH\H~\OH o.oe m4Ho s.mm H.O ooHno nuo\oouounos\omn .ooo~\mH\uooHoem om om OH OH soHuHosoo ooom\soHuoHuo<\soHusHom HmsumOHHmsu m mo mCmsEv .mossm sou mCHBOm usumm msumo v um COHumCHEHso ossm quousm \xnsooc smme HoomxHos~ same room quEumsuB ossm A.o.uCOOV .OH mqmfifi xHQmem4 1293 02615 902