TRAN3L0CATION STUDIES IN THE SUBMERGED TISSUES OF AQUATIC VASCULAR PLANTS by Eugene T. Oborn AN ABSTRACT Submitted to the School of Graduate Studies of Michigan State College in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of Botany Year Approv red () 1f / • r/( /9 s f 19f?l In order to grow diversified crops on much of the land in the western United States i t is necessary to supplant the moisture provided by nature in the form of rain, snow, e tc ., with additional water which reaches farm lands through established irrigation canal distribution systems. These systems frequently support heavy growths of vascular aquatic plants which prevent or slow down the passage of water. Since reducing the carrying capacity of a canal makes i t necessary to deprive some potentially crop-producing land of the required water to bring the crop to a satisfactory harvest, i t is imperative to keep the water­ ways open. This thesis is a report of certain pertinent investigations which suggest improved and more effective field techniques to accomplish a solution of the problem a t hand. Broad- and narrow-leaved c a tta il, water sedge, true waterweed, American pondweed, horned pondwreed, leafy pondweed, Richardson*s pondweed, gigantic sage pondweed, and slender sago pondweed plants were transplanted from the field directly to containers with as l i t t l e root disturbance as possible. Spraying of the submerged aquatic plant materials in the water drained tanks was performed with a 1 -quart capacity model A, Sure Shot pneumatic sprayer. Lethal effects of 2,1^-D ester appeared to be transmitted through the immersed c a tta il leaf, past the waterline, and into the crown of the plant. No shoot regrowth developed in ester treated plants. When broad— leaved c a tta il roots were immersed for 21; hours in 10 ppm of the s a lt and ester forms of 2 ,U-D, and the ester form of O born— 1 2 ,l 4 , 5>-TJ e f f e c t o f t h e p a s s a g e o f t h e s y s te m ic h e r b i c i d e s i n t o t h e p l a n t s was e v id e n c e d b y t h e f a c t t h a t no s h o o t r e g r o w th w as i n e v id e n c e 8 w eeks f o llo w in g l e a f h a r v e s t . When aerial herbicidal treatments were made the following single or repeated 2,U-D applications were effective in obtaining complete, or nearly complete, eradication of the waterweeds growing in the soil bottom of the treated tanks. Name of plant Narrow-leaved ca tta il Broad-leaved c a tta il Water sedge True waterweed Leafy pondweed American pondweed Gigantic sago pondweed Richardson1s pondweed Horned pondweed Pounds per acre Number of treatments Percent eradication 25 1 1 1 1 1 1 2 2 2 100 100 85 100 100 95 95 98 100 m 27 5 5 10 20 12 11 A study was made of the changes in ca tta il root reserves in under­ ground plant parts which are due to seasonal growth phenomena. Weekly measurements were made throughout the entire growing season in an attempt to correlate below-ground carbohydrate root reserves with easily observed above-ground phenomena. Broad- and narrow-leaved c a tta il roots showed considerable variation in the amount of carbohydrates present during the growing season. Highest carbohydrate was present during the winter dormancy period and lowest carbohydrate was associated with production and maturation of male and female fruiting bodies. Root reserves are low from the time of the f i r s t appearance of the fruiting stalks until pollination has been completed. Oborn—2 In the narrow-leaved and broad-leaved c a tta il growth s ite s , not inundated by water, root reserves were a t a minimum when the plants had attained a height of appro:cLmately 100-130 cm and 5>0-120 cm above the ground line respectively. O b o rn — 3 TRANSLOCATION STUDIES IN THE SUBMERGED TISSUES OF AQUATIC VASCULAR PLANTS A Thesis Presented to The Guidance Committee Michigan State College In P a rtia l Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School Department of Botany Ly Eugene Timbrell Oborn May 1951 ACKNOW LEDGM ENT Sincere appreciation is expressed to Dr. B. H. Grigsby, Associate Professor of Botany and Research Associate, who directed the work here reported. Acknowledgment is also due the United States Department of Agriculture, Bureau of Plant Industry, Soils and Agricultural Engineering, and the United States Department of the Interior, Bureau of Recla­ mation, for the equipment and fa c ilitie s generously made available to carry out this study. CONTENTS Page Introduction ..................................................................................................................................... Anatomy of Aquatic Vascular Plants .............................................................................. Seasonal Carbohydrate Root Reserve Trends in Roots and Rhizomes of Broad- and Narrow-Leaved Cattails .......................................................................................................................................... 1 5 7 Date of collection . . . . . . . 11 Notes pertaining to appearance of fruiting bodies, pollination time, etc.............................. . . . . . . 12 Weekly water temperatures approximately eight inches belovf lake s u r f a c e .....................................................................................13 Height of plants above water surface or ground line . . . . . 13 Total length of plant shoots ..............................11+ Total length of seed s t a l k .................................................................... 11+ Length of female and male s p ik e s ................................................................... . 11+ Width of female s p i k e ............................ V~> Width of leaf ........................................................ 15 Shoot and root fresh weight ............................................................15 Percentage dry matter in shoots and r o o t s ...................................................16 Percentage carbohydrate in oven-dryroots ......................................16 Identification of S p e c ie s ....................................................................................................19 Materials and Methods . . . . . .................................................................................. 21 I. II. III. IV. V. Leaf-Tip Immersion Test ................................. 21 Root-Immersion T e s t ..................................................................................22 Aerial Herbicidal Treatment onPlant Materials . Transplanted to Buckets 3-Gallon Capacity . 23 Aerial Herbicidal Treatment onPlant Materials Transplanted to Metallic Containers of 27-l/2-C-allon C a p a c ity .....................................................................28 Miscellaneous Tests ...................................... 32 Discussion of R e s u l t s .................................................................... I. II. III. 31+ Leaf-Tip Immersion T e s t ...............................................................................3U Root-Immersion T e s t.........................................................................................35 Aerial Herbicidal Treatment on Plant Materials Transplanted to Buckets of 5-Gallon Capacity............................................ 36 ggge IV. Aerial Herbicidal Treatment, on Plant Materials Transplanted to Metallic Containers of 27— l / 2 -Gallon C apacity..................................... True waterweed........................................................................................... Leafy pond-weed ..................................... American pondw eed................................................................................ Gigantic sago pondweed ........................... Slender sago p on dw eed.................................................... Richardson* s pondweed ........................... Horned pondweed . . . . . . . . . . . . . . . . . V. Miscellaneous T e s t s ................................ Detection of 2 ,U-D in roots of toptreated c a t t a i l plants .......................... Effect of systemic herbicidal applications on water sedge ............................................... 38 39 39 39 [j.0 ill i|l I4I kk UU ij.it U6 Summary .......................... Literature Cited Appendix ii INTRODUCTION For the past several years, the number of acres of land placed under irrig a tio n in the western United States has increased. This same land, without additional water to supplant th a t provided by nature in the form of ra in , snow, e tc ,, has been to ta lly without, or of much le s s , value from the standpoint of growing crops. To provide th is water, not only is i t necessary for riv er waters to be impounded and irrig a tio n canal d istrib u tio n systems established, but also for the irrig a tio n waterways to be free of obstructing plant growths which prevent or slow down the passage of water. Reduction in carrying capacity of a canal can mean only one thing; i . e . , some p o ten tially crop-producing land w ill be deprived of the necessary water to bring the crop to a satisfactory harvest. Therefore, i t i s imperative to keep the waterways open. Balcom ( l ) , Crafts (5), and Speirs (33) > have discussed the several methods used in the past in the attempt to keep the waterways open. Among these are chaining, draining, dredging, burning, bio­ logical control, and chemical control. None of these have been satisfactory or generally permanent because the roots, rhizomes, or other propagules of the treated aquatic vascular plants have not been k ille d . Several chemicals, separately and in various combinations, have produced brovming and defoliation of waterweed leaves, or a temporary k i l l of only the top portion of the plant in irrig a tio n ditches but the viable rootstocks have remained as a source of new growth. Thus, i t i s not only desirable to destroy vegetative growth pres­ ent in an irrig a tio n channel, but also to alio?/ an absolute minimum of fru its or seeds, la te r a l or winter buds, or fragmentation propagules to remain behind for new in festatio n . At the beginning of each growing season many vascular plants use the starch food reserves stored in th eir roots to build new plant tissue. This process continues u n til such a time as the chlorophyll- bearing tissue of the new plant can manufacture food in the quantities needed for the new growth. As plant growth slows up, the chlorophyll- bearing tissue in the new plant is able to catch up with, and then surpass, the immediate demands for food used in the manufacture of new tissue^ this re su lts , of course, in starch reserves being put back in the underground storage organs for use a t a la te r date. Chemical analyses, presented la te r , of the broad—and narrow­ leaved c a tta il rootstocks show that carbohydrate is the principal form of food reserve storage. I t seems probable that the other aquatic plants discussed in this paper store th eir food reserves in a like manner. Evans, Mitchell and Heinen (7)> Rasmussen (25) s Smith, Hamner and Carlson ( 3 2 ) , have shown that phenoxyacetic acid derivatives, in herbicidal concentrations, stimulate plant respiration, food digestion, and u tiliz a tio n of reserve food materials. Freeland (12) observed that 2 ,U— D a t concentrations of 30 and 100 ppm decreased the rate of photosynthesis in true 'waterweed (Anacharis 2 canadensis) , -with the higher concentration being more effective. Mitchell and Brown (20) treated morning— glory (Ipomoea lacunosa) plants with 2 ,U— B in herbicidal concentrations and. found a depletion of readily available carbohydrates within a 3 -week period afte r treatment. Hall and Hess (lU) have suggested th a t the re la tiv e ly high tolerance of submerged aquatics to 2 , 1*— B i s associated with th e ir reduced vascular systems and resulting lim ited transport of phenoxyacetic acid derivatives. Braining water off of the plants before treatment would therefore appear essential for the most effective con- . tr o l of submerged species with 2 , 1*— B. So fa r as i s known, Surber, Minarik and Ennis (31*) are the only workers who have controlled normally submerged aquatic plants by f i r s t draining off the water and then spraying with systemic herbicides, the flaccid , rooted aquatic plants lying on the s o il bottom. In th is case, a pond supporting American pondweed (Potamogeton nodosus) was drained and bottom sprayed with 2,1*-B dissolved in tributylphosphate a t the ra te of 10 pounds 2 ,L*— B acid equivalent per acre. American pondweed shoots faile d to reappear a fte r th is herbicidal treatment. I t w ill be shown th at when broad—and narrow— leaved c a tta ils (Typha l a t i f o l i a and Typha angustifolia. respectively) and water sedge (Garex aquatilis) were treated with phenoxyacetic acid derivatives in herbicidal concentrations, a depletion of readily available carbo­ hydrates occurred in the underground roots and rhizomes of these plants. Plant regenerative power is at i t s lowest ebb when food reserves 3 are low. Accordingly, application of any given herbicide, contact or systemic, at the time when food reserves are low should effect the most successful eradication or control with expenditure of minimum time, effo rt, and cost. Because of recent widespread success of certain phenoxyacetic acid derivatives on landweeds, the effect of these compounds on obnoxious submerged aquatic weeds was believed to be worth investi­ gating . To th is end, translocation studies in the water-submerged tissues of aquatic vascular plants were made. Since carbohydrates in the crown, rhizome, and root of the c a tta il plants appear to be equally readily available to the plant for new growth, the word "root" is used in this study to include that part of the underground tissue below the line from which fibrous roots emerge. h .ANATOMY OF AQUATIC VASCULAR PLANTS Structural changes in vascular hydrophytes which Eames and MacDaniels ( 6 ) considered as adaptations to aquatic environments are reduction of protecting, supporting, and conducting tissues and frequent provision for aeration the f u ll length of the plant, i . e . , leaf, stem, root, by the development of air chambers. Finely divided, terete leaves allow for increased contact between leaf surface and surrounding water. Absorption of gases and nutrients takes place directly from the water by stem and leaf • The epidermis commonly contains chloroplasts and may thus form a considerable part of the photo synthetic tissu e, especially where the leaves are very thin. In submerged tissues of hydrophytes, stomata are wanting. The floating leaves, however, e .g ., in American pondweed (Potamogeton nodosus), have abundant stomata upon the upper surface. The greatest proportional reduction in the vascular tissue occurs in the xylem (see plates IX and X) . In many forms the xylem consists of only a few elements, even in the stele and main vascular bundles. xylem elements are entirely lacking. Less commonly, In these cases there is usually a more or less well-defined xylem lacuna to mark the normal position of the xylem. Phloem tissues, on the other hand, are in most cases fa irly well developed as compared with the xylem. They resemble the phloem tissue of reduced herbaceous plants generally, in that the sieve tubes are small as compared with those of woody plants. 5 The endodermxs is often weakly developed or may be entirely wanting, I t is important to note here that Mitchell and Brown (20) found that the stimulus resulting from 2,U*-«D application on bean plants was closely associated with translocation of organic food materials and that this stimulus occurred as a continual flow under conditions favorable for carbohydrate translocation and probably was confined to the living phloem or parenchyma c e lls , Thimann (3£), Weaver and DeRose ( 3 6 ) have shown that stomata are not important portals of entry for phenoxyacetic acid derivatives and that, when applied'to the aerial portions of plants, the compound moves downward whenever translocation of synthesised materials occurs. This downward movement of the phenoxyacetic acid stimulus through the living phloem or parenchyma cells is very important for, as pointed out, the phloem tissues of aquatic plants are in most cases well developed as compared with the xylem. 6 SEASONAL CARBOHTBRATE ROOT RESERVE TRENDS IN ROOTS AND RHIZOMES OF BROAD- AND NARROW -LEAVED CATTAILS I t w ill be pointed out further on in this study that i t is possible for as many as 1 5 -2 0 new plant en tities to emerge from basally clipped c a tta il rootstocks. Apparently so long as la te ra l buds and adequate carbohydrate and other food reserves are retained in the underground parts, these clipped plants, other conditions being favorable, are capable of regenerating new plant growth. I t was desirable, accordingly, to study the changes in root reserves in these underground plant parts which were due to seasonal growth phenomena. Having established the period, or periods, of low root reserve i t would thus be possible, by making herbicidal appli­ cations when root reserves are at a low ebb, to get maximum herbicidal effect with minimum time, effort, and cost. As w ill be shown la te r , the majority of root reserves in broadand narrow-leaved c a tta il were found to be in the form of carbohydrates • Protein and crude f a t determinations were made on broad- and narrow— leaved c a tta il root samples in accordance with methods suggested by Lepper (17), Loomis and Shull ( 1 8 ), and Woodman (37) • Root samples of both species were taken from water inundated growing sites on April 1 0 , 1950. In th is study of seasonal carbohydrate root reserve trends on broad- and narrow-leaved c a tta il roots and rhizomes (see plates IV and V), weekly measurements were made throughout the entire growing season in an attempt to correlate below— ground carbohydrate root reserves with easily observed above-ground phenomena. C attail carbohydrate root reserves were determined on the samples taken a t v/eekly intervals and associated above-ground phenomena were recorded for the following situations: 1. 2. Growing s ite inundated by water a. Fruiting broad— leaved c a tta il b. Nonfruiting broad-leaved c a tta il c. Mixture of fruiting and nonfruiting narrow-leaved c a tta il Growing s ite not inundated by water (subirrigation) a. Mixture of fruiting and nonfruiting broad— leaved c a tta il b. Nonfruiting narrow-leaved c a tta il Weekly notes on the above-ground phenomena for c a tta ils in the above five categories were recorded as fol3.ows: 1. Date of collection 2. Appearance of fruiting bodies, pollination time, etc. 3. Weekly water temperatures approximately eight inches below lake surface Uo Height of plants above water surface, or ground line 5>. Total length of plant shoots 6. Total length of seed stalk 7. Length of the female and male spikes 8. Width of female spike 8 9• Width of leaf Weekly laboratory data were recorded as follows: 1. Shoot and root fresh weight 2. Shoot and root oven-dry weight 3. Percent dry matter in shoots and roots U. Percentage readily hydrolizable carbohydrate in ovendry roots After field linear measurements had been made on about seven specimens from each grouping as listed earlier, the cattails were dug, brought into the laboratory, the soil washed off the roots, and the remaining laboratory linear measurements made. The roots were then separated from the shoots and cut up into approximately l / 2 -inch lengths (line of separation determined by the point of emergence of fibrous ro o ts). The shoots and roots were placed in separate paper sacks and weighed. The shoots and roots were then placed in separate six-inch cubical wire baskets and heated in a drying oven at 100° C for 30 minutes as suggested by Miller (19) to inactivate the enzymes present. The oven temperature was then lowered to 62° C and the plant material heated for 72 hours. The oven-dried shoots and roots were transferred to separate paper sacks and reweighed. The shoots were then discarded and the roots were stored in paper sacks in a cool, dry place so that later chemical analyses could be made on the specimens collected at weekly intervals through the growing season. Carbohydrate root reserve in broad- and narrow-leaved c attail samples was determined as follows: 9 Preparation of sample. The dried, stored roots were pulverized in a Quaker City mill so that approximately 99.8 percent passed a lU-mesh sieve. Table 1 shows the sieve analyses on 10 of the grinds used. The pulverised sample was placed over a nest of the Tyler standard screens of the sizes indicated, placed in a Ward shaker, and shaken for a period of five minutes. To insure a uniform sample being taken, a ll the ground sample passing the No. 16 sieve was thox’oughly nixed with a spatula after which a three-gram sample was weighed on a torsion balance. I t was recognised th at, in c a tta il roots, starch is accompanied by root reserves such as pentosans and various hemicelluloses that also yield reducing sugars upon hydrolysis. The method of direct acid hydrolysis was relatively quick and easy of execution, and sufficiently accurate to give results comparable with published analyses. 10 TABLE 1 TYLER STANDARD SCREEN SIEVE ANALYSES ON REPRESENTATIVE PULVERIZED CATTAIL ROOTS SIEVED FOR 5 MINUTES IN A WARD SHAKER S i e v e Number M eshes -bo t h e In c h Sample : T o ta l Number : : Grams : 2 3 . 8 1 :P e r c e n t: 1 0 0 .0 0 : Grams : 2 9 .7 2 :P er ce n t: 1 0 0 .0 0 : Grams : 2 3 .1 3 :P er ce n t: 9 9 .9 9 : Grams : 1 9 .6 5 : P er ce n t: 1 0 0 . 0 1 : Grams : 1 7 .3 5 : P e r c e n t: 9 9 .9 9 : Grams : 1 7 . 2 ...... :P e r c e n t: 9 9 .9 9 : Grains : 2 5 . 0 7 :P e r c e n t: 1 0 0 .0 0 : Grams : 3 h .$ 8 :P e r c e n t: 1 0 0 .0 0 os : Grams : 3 3 .0 : P e r c e n t : 1 00.00 : Grams : 3 9 .2 10 : P e r c e n t : 100.00 A v e ra g e p e r c e n t: 16 11; 30 28 0.0 0.6 0.0 2 .5 2 0 .3 0.1 0 .5 3 0 .0 0 .1 0 .5 7 0.1 0 .5 8 0 .0 0 .0 0 .1 0 .3 0 0 .0 0 .1 9 5o 58 100 100 5 .5 22.69 5 .3 1 7 .8 5 5 .5 1 9 .5 8 2.9 6.8 200 20 0 Pan 5 .0 6 .0 2 5.21 28.57. 7 .0 9 .0 6 .1 3 0.3 0 1 .01 2 3 .5 7 . .27*27. 6 . 6 5 .6 7 .0 0 .3 1.3 0 28.57. 1 9_ .9 !. . 3 0 ,3 0 . 8.0 0 .2 5 .8 3 .7 5 0 .8 2 18.88 1.02 111 . 8 0 25.59. 5 .8 3 .6 5 .6 0 .3 3 .1 1 .7 1 17.71_.__ 3 2 . 0 0 . 20 .5 7 . . 27-kL. 5 .8 0.2 5 .7 3 .9 3 .5 1.1 6 22.67 ..2,1.32 20.3 5.... 2 7 .9 1 0.1 5 .3 6 .3 5 .5 8 .9 2 1.20 2 5 .2 0 3 5 .6 0 o.Uo 1 7 . 6 0 8 .1 0 .1 12.7 5 .2 8 .5 36.81 2 3.5 8 0 .2 9 1 5 . 0 7 2_5*35_ 9 .8 7 .8 6 .9 7.7 0 .7 2 9.70 2 .1 2 2 0 .9 1 23..6k 2 3 . 33 . 1 5 .0 8 .2 1 0 .0 o .6 6 .5 20.92 i . 5 3 _ 16.33. . .2 5 , 5.1 2£ J i 28.85 1 8 .2 8 21.99 29.39 21.01 Results of protein, crude f a t , and carbohydrate determinations made on oven-dried residue of broad- and narrow-leaved c a tta il root samples, both taken from a. -water inundated growing s ite on April 10, 19E > 0 , are presented in Table 2. TABLE 2 CRUDE FAT, PROTEIN, AND CARBOHYDRATE CONTENT OF CATTAIL ROOTS Name of plant Percentage Crude Fat**- Narrow-leaved c a tta il Typha angustifolia L. 0 .£ 2 8 0 Broad— leaved c a tta il Typha l a t i f o l i a L. 1.2933 1. 2. 3. Percentage Protein 2 Percentage Carbohydrate^ 60.17 6.73 SU.li.0 Lepper (17) 9 Woodman (37) Lepper (17) s Woodman (37) Loomis and Shull (28), Schaffer and Hartmann (31), Woodman (37) From th is table i t can be seen th at re lativ ely small quantities of crude f a t and protein are present in broad—and narrow— leaved c a tta il rhizome and roots and th at the root reserve i s primarily in the form of carbohydrates. Results from chemical analyses of c a tta il root and rhizome samples collected a t weekly intervals throughoivt the growing season are pre­ sented in Tables 3~7. These data show the changes in carbohydrate materials (see plate V) in these underground plant parts which are due to seasonal growth phenomena. Date of Collection Weekly c a tta il root collections and growth measurements began 11 TABLE 3 FIELD CARBOHYDRATE ROOT RESERVE STUDY OK MIXTURE OF FRUITING AND NONFRUITING NARROW—LEAVED CATTAIL PLANTS DEVELOPING FROM GROWING SITE INUNDATED BY WATER Specimen Number Date 3- 20-50 3-27-50 U-3-50 U-10-50 h I 4- 1 7 - 5 0 5 6 U— 2 J4.— 5o rj cl 5-1-5.0 8 5- 8-50 . 5- 15-50 9 10 5-22-50 lib 5- 29-50 12° 6-5-50 6-12-50 ......13 . 6-19-50 lUd 6-26-50 IS ? 16* .7-3-50 7-10-50 _ . 17 . 18 7-17-50 19 7-2U-50 20 7-31-50 21 8-7-5o 22 8-1U-50 8-21-50 . 23 8-28-50 2h .9-5-50 25 26 .9-11-50 9-16-50 27S 28 . 9-25-59 10-2-50 .... .29 -19.. , 1 0 - 9 - 5 0 1 2 3 . .. . a b c d e f Height Lake Surface above h2o h2o Surface Temp op cm U5.8 51.8 53.1 5U.2 52.2 57.5 . 51.2 52.5 _ 66.0 76.5 60.1 7U.0 73.0 75.0 8U.0 82.5 80.5 71.5 7 0 .0 72.5 78.5 79.5 66.3 53 .0 . 76.5 57.0 6 1 .6 60.5 5 5 . 0. 5u.o Final Length cm Final Width cm Seed Female Male Female Leaf Stalk Spilee Spike Spilee Leaf 8.9 10.5 33..8... 2U.1 k7.3 68.9 10^.7 .91.0.. 107.3 125.9. 107.6 130.1 130.5 1U2.5 160.h 163.2 1U9.5 163.3 172.3 177.3 180.7 18U.2 17 .8 . 9 ...20^ 27-9 33.7 . 38,9. 6 3.1 76.3 93.8 8L.3 115.9 150.7 130.2 133.9 11+8.2 1U2.U 151.1 178,3 .157,6 1 6 8 . 1+ 188.6 2o"5.6 197.9 191.9 206.8 162.8 I06.2 1 8 0 .1 18I+.2 217.8 229.0 172.5 106.9 0.6 0 .7 0.6 119.6 12. U 19 .k . 0 . 5 . 11.6 0.6 128.0 10.9 10.6 128.7 13.5 0.9 m i . 5 10.6 1U.9 0.9 13U.U 10.9 .. I k . 9 1.2 1.6 11.7 11.3 133.1 I.®7 l5.2.£ 13.7. 1.7 10.9 123.9 1 0 . 8 1.8 1 5 2 . 0 .. 13.3 12.7 152.7 1 *T 1.8 iL ..I. 162.5 1.8 1 SU. 1 16.5 1 .8 1 7 2 . h 17,)( 1 6 9 . 0 _17.1 1.7 167.6 1 5 .6 1 .8 1 .8 1 7 6 . 1. _.17,3. 1 6 .8 1 .6 1 72.3 1 .6 159.2 i5.U Cattail -tops frozen back 6 to 8 inches. Green c a tta il leaves knocked down to water surface by snow, Narrow-leaved fruiting bodies beginning to appear, Narrow-leaved c a tta il beginning pollination, Narrow-leaved c a tta il pollination 50 percent completed, Narrow-leaved c a tta il pollination completed. Female spike cinnamon brown. °-7 0 .6 0 .5 0.6 0 .6 0.6 0 .7 0.8 0 .7 0 .8 0 .7 0 .8 0 .6 "o.B 0.9 0.9 0 .8 1 .0 0 .8 Percent Carbo­ hydrate Specimen Fresh g : Ovan-Dry g OvenNtunber • • dry Shoots Roots: Shoots Roots Shoots Roots Roots 110.8 52l*«6: 21.1 1 _.5&,53 . 2 6 .1 1 5 ,2 2 690.1: 1.9 63.33 180.3. . 12,5 2 6 .6 65.77 . . . 3. 3.7 5l*3.2: 0.5 lfh-7 ,13_.5. 1.7 16.2 178.8: 60.17 1*9.3 . 1 0 , 5 .. 22.0 h 62.6 1*50.1: 5.6 I 2 8 .I* 9.0 6 S.1 3 28.5 .5 8.8 6 81*0.3: 15.6 . 17.7.2 1 9 9 .0 23.7_ - 6A - 7 2 llf.2.6 678.5: 15.1* 10.8 23.2 1 5 7 .7 6 3 .* 7 7 8 2 2 . 1* .331.2 8 5 8 . 0 : 29.5 1 9 1 .9 8.9 6 6 .0 3 6 6 .9 0 9 5p5..7 803.U: 52.1* . 1 7 9 , 6 . 1 0 . 1* 2 2 . 1* 61*5«2 9 9 1 . 9 : 8 6 .8 21*5 . 0 10 21 *.7 5 9 .0 3 _ 13.5 6 0 .9 0 11 56.1 1 9 .2 21*1*.7 _ 292.3J 32,5.. 13.3 3 0 1 . 1* 21*1*.0 : 52.2 12 U2.97 . 1*5.0 17,3. . 18.1* 2 2 .2 6 2 .I* 525.1 .396,3:13.7.6 i a .7 7 13 . . . . . . . 15.7 91*.8 .. 15,5. . 17.7 536.1*: 78.0 it* . 59.27. 1*8 0 . 1* 573.U:109.3 18.8 22.7 15 . . 108.3 5 6 .2 3 16 U6 .0 0 5oo.o 35.U,5: 8 5 .8 1*3 .2 1 2 .2 17,2 2 2 .6 17 377. U 50U.3:130.6 79,3 _15,7_._ *9.03. 18 L *0 l*.i*: 9 2 . 1 6 i.b 15.2 4 6 6 .3 18.9 U3.93 lifi.90 1 2 .2 17.2 7 3 U .7:132.1* 89.1 19 7S6.3 20 1*8 5 .8 316.8:108.8 2 2 . 1 * 53..30 . 16.5 85.3 21 27.1 §6 0 . £. 1*18.3:1^1.8 17.3 72.5 57.03 22 27,6 530.7 1*01.9:11*6.5 ... .7.1,9 17.9 58.17.. 5 8 .8 0 23 ^76.3 . 535.1:11-13,7 15.9 21 *.9 85-3 2 7 .8 2 0 .6 l*6l*.o 3 0 2 . 0 : 1 3 1 .6 2h 82.3 63.3327.6 2 0 .6 2 l* 6 *. 03 J 1 3 .8 1*73.6:11*3.1 98.7 26 21*.2 1 7 .8 .5.23.8 291.1*: 126.7 6 l.5 o 51.8 59.90 611.6 382.2:165 .9 20.0 27 27.1 76.5 28 3 6 ]*. 1* 391.5: 92.5 67.23.. 25,U 20.5 80.3 6 i.5 o . 29 59.3 17.8 365.7 335.9: 78.5 21,5 6 * .io 2l*. 2 25.6 3 2 2 .8 2 1 1 . 8 : 82.6 51.2 3P. . —....... Final Weight Percent Dry Matter . H • on o *A . .. . _ . g Considerable yellowing in c a tta ils TABLE U FIELD CAEBOHZieATE ROOT RESERVE STUDY ON NONFRUITING NARROW— LEAVED CATTAIL PLANTS DEVELOPING FROM GROWING SIT E NOT INUN­ DATED BY WATER Specimen Number ... 1 2 3 h a b c 3-20-50 3-27-50 U-3-50 L-lo-50 U-17-5o .U-2U-50 5-1-50 5-8-50 5-15-50 5-22-50 5-29-50 6-5-50 6-12-50 6-19-5o 6— 26— 50 .7-3-50 0 ir\ 1 0 H 1 C “- 5 6 7a 8 . 9_ 10 lib 12 13 1U . 15. 16 17 18 19 20 21 22 23 2U . . . 25 26 27° 28 29 30 . Date 7-17-50 .7-21-50 7-31-50 .0-7-50 6-iU-5o 6-21-50 8-28-50 9-5-50 .9-11-50 9-18-50 9-25-50 10-2-50 10-9-50 Lake Height Final Length :Final Width Surface above cm : cm h2o Ground Temp Surface : Seed Female:Male :Female: op cm Leaf: Stalk Spike : Spik e: Spike :Leaf U5.8 5 1 .8 53.1 5U.2 52.2 U.7: 57_.5 5U.2 52_.5 6 6 .0 : : : O.k 23.9 . . 23.3s . 76.5 _. 2 8 .8 6 6 01 31.9: 7A.0 : : :0 .6 . 35.0 _ L2 . 8 : 5 6 .1 : 73...0 : : : 0 .5 . 5U.U 7 2 .6 : 75-0 . 6 7 .9 ...............:.. t :0.7 81)..0 : : :0.7 87.9 86.3: 105.6: : : :0.7 §2 .6 101.7 : : : 0 .8 121.8 : 80.5 120.3 1 2 6 .1 12R.5: : : :0.9. .71.5 : : :0.7 70.0 1 2 6 .u 111.5: : : :0 .8 136 .L 135.0: 72.6 : : :0.8 luO.L: 78.5 lLl.5 : : :0.8 1L2.9: 1U2.7. 79.5. : : :0.6 1L0.9: 66.3 1 3 6 .3 53..o_ 1L8.5 : : :0.7_ 1U6.7: : : :0.8 152 ..8 : ... lL o.l 76.5_ I l 6 .6 : : : :0.7 57.0 138.5 6 l.6 : : :0.9 150.7: 1 3 8 .3 1 1 1 .1 : t : :0.7 1 3 7 »3_ 60.5 : : :0.8 55.o 1U2.2 130.9.: . _ . 5k.o l-23.i4-_._ 1 1 6 .1 : * * *0-7 — C attail tops frozen back 6 to 0 inches, Green c a tta il leaves knocked down to water surface by snow, Considerable yellowing in c a tta ils . Percent Percent Carbo­ hydrate Specimen Dry Matter Oven-Dry g OvenNumber Fresh g dry Shoots Roots Shoots Roots Shoots Roots Roots 1 26.5 62.27 91.7 316.5. 65.0 2 6 .2 2 57.10 2U7.9 27.0 213.0 57.It 4 1 .3 7 ....... 3 27.2 U2.6 it i56.lt 59.^3 19.6 86 .it 22.7 . 51.70 5 6 2 8 .6 22.0 ... 53.23 129.8 21.0 it8.03 7 121.7 25.J 8 itit.itO 123.8 20.It l6.5 19.8 37.8 U9,53 9 190.7 57.8o 10 lit.8 1 0 6 .5 U79.0 15.8 112.0 23.it 11 21.2 . 56.67 1 6 9 .7 769.0 22.5 162.9 13_.3 12 1 U.0 105.5 87.8 U6U.7 22.7 5 1 .7 7 15.9 13.8 21.2 128.6 29.6 13 21^ .0 £0 6 .0 .. 53-t3_ lit 52.8 u9 .93 182.1). 309.it 2U.9 13.7 17.1 23.0 21.1 15 57.97_ 265.3 223.1 6o.3 51.3 16 13.8 18.8 210.it 111.2 U 3.63 39 .6 l5 X 17 UU.8 21.1 .17.5 h i.h i 85.7 U05.3 256.0 18 lit.6 . U3.00 207.8 UO8 .8 72.It 30.it 17.7 20.6 19 it8U.it 881 .i-f 100.0 138.6 15..7 . h f . 0 1 . 20 5U .80 18.8 2 6 .2 3U0.2 20U.0 89.1 36.3 21 32.8 29.0 55.77 272.0 168.9 78.9 22 it9 .2 0 2 6 .1 1 6 . It 2 6 .3 3 1 6 . 7 l 6 o . & ~ 82 T0 23 2 2 .0 50.8 8 0 .0 30.7 5U.9_7. 293.1 2 3 0 . 6 2 it 2 2 .8 3 0 .8 6 1 _.it 2 U1 .8 279.3 63.7 7U.5 25 2 6 .1 3 2 .8 6 0 .0 3 6 2 j t '" t i k i 190.9 179.5 26 59.90 2 3 .6 31.3 19li.3 2“SoT.T 6 1 . 2 ' " 66 0% 27 U2 .1 63.P2. 28.7 277.8 1 9 1 .0 1 1 7 .0 5U.9 28 6 1 .1 0 29 .it 33-1 65.7 179.2 223.U 59 .It 29 61.23 2M 29.3 6 it.Lt 312.6 259.3 92.1 30 6 2 .2 0 u5.o 2it.7 52.8 3 9 .7 . 117.3 l6o.9 Final Weight. _ ............ . . . .TABLE £ .FIELD CARBOHYDRATE ROOT RESERVE STUDY ON NONFRUITING BROAD­ LEAVED CATTAIL PLANTS DEVELOPING FROM GROWING SITE INUNDATED BY WATER Specimen Number Date HoO 1 2 . 3 U 3 . ... 6 7d 8 . 9. _ _ 10 llb 12 13 . .. I k . . 1? 16 . 17 ... . 18° 19 20 21 22 23 2U _.... 25. 26 27a . 28 29 30 a. b c d . .. Lake Height Surface above Final Length cm Final Width cm HoO Temp : Surface °F : cm 3-20-50_ U5.8 : 1 0 . 2 3-27-50 5.1.8 . : 1 2.7 : liuO Jte3=£o & ..1 U-1 0 - 5 0 i k . l _ : 27.3 U-17-50 £ 2 . 2 : 2 U. 8 : U9 oh _U-2 i r f o 57 5-1=50 5li.2 :. . 53...U. 5-8-50. 5 2.5 . : 78.9 6 6.0 5r.l^r50 : 100.7 5-22-50 76.5 : 137.2 : 129.2 5 - 2 9r50 6 6 . 1 6-5-50.. 7U.0 : l U l . l 6 - 1 2 - 5 0 .73 . 0 : 13U-3 6 - 1 9 - 5 0 7 5 .0 : J 3 S -7 6 - 2 6 - 5 0 8 U. 0 : 178.5 7-3-50 82.5 . : 180.7 7 - 1 0 - 5 0 . 80.5 . 177.0 7-17-50 71.5 17.3*6 7-2U-50 70.0 17 -6 . 8 7-31-50 . 12,5... : 182.9 8-7-f>0 7.6.5.. : i s 6 . u 8-1U-50 79 .5 . t 190.6 8-21-50 6 6 . 3 : 2 0 8 . 6 8 - 2 8 - 5 0 5 3 .0 . : 192.8 9 - 5- 5o 76.5 207 .U 9-11-50 5.7.0 . : 199.6 9-18-50 6 1 . 6 t 195.8 9 -25-50 6 0 . 1 : 196.55 5 .0 10-2-50 169.-.U 10-9-50 -,5 k -P : 1E>8.0 Seed Female:Male F emale Leaf S ta lk Spike : Spike Spike Leaf 20.8 28.5 2 9.7 28.2 62.7 69.5 82.7 103.3 130.1 1U5.0 11+7 . 3 l5 ? -5 156.2 163.8 188.9 2 0 9 .5 185.3 191 .1 210.3 180.8 213.1 221.7 21x0.9 229.0 219 .1 2U7.2 21+3-5 232.6 235.0 208.u Cattail tops frozen back 6 to 8 inches Green ca tta il leaves knocked.down to water surface by snow, Broad-leaved ca tta il vegetative tips browned, Considerable yellowing in c a tta ils. 1.0 1.0 i-5 1-5 1.6 1-5. 1 .7 _ 1 .6 1-5 l.U l.U l.S 1-5 1 .3 1.5 l.U i-5 1.5 l .U 1-5 1-5 l.U 1 . 1+ 1.2 P ercent Carbo­ hydrate Specim en O venNumber F r e sh g : Oven-Dry g • dry S h oots R o o ts :S h o o ts R o o ts S h o o ts: R o o ts R o o ts 2 6 .6 55.10 1 8 .1 1 2 6 .6 17U.3t 6.3: l 6 .k Ik . 2 51.2? 2 1 ^ .2 6.9: 62.3 1 * 1 * 0 .3: 220.9 U 6 .8 362.9:. 8.3: lg -2 _52_.83 ^•9 ~ §1 ,1 3 . 51*.1*0 90.7 . 7.9: 16.3 l 6 ^„2 555.6.: 13.1 1* 8 . 6 : 50.77 9.1 5 .... .... . 1 0 ^ .2 217.8: 29*3 13-5 6 7 . 0 : 15.1 39.77 553.6 299.2: 3 8 .8 . k§ ,2 8 . 1*: il*.i* .... 1*8.53 7 _____________ 93.9 56k.9 6 5 2 . 0 : k7.5 8 7 .0: U7.1 39.-8 673.3 ia o .i: 9.-7 . 38 .J33 2 8 .6 0 8 .2 : 9 8 «§ k7.7 1325.7 559 .k: 107.8 10 29.70 1 0 .1 9 . 6 : 1 0 6 7 .2 918.3: 1 0 2 .2 92.9 11 25.20 9 .8 : 7.9 1*8.9 1019.1 618.9: 100.0 12 1 2 .1 : 29.77 6 l*.l* 91*6.7 666.3: Ilk .6 9 -7 2 2 .1 0 1 1 .2 lk . 8 : io .i 13 660.7 U51.8: 9 8 .0 1 0 . 8 : 11* 27.27 3 6 .3 128.8 7 . 1 5 0 8 . 0 : 1195.9 31.70 1 6 . 6 : 6 1 * .8 9.§ 15 _________ . 835.6 6 8 3 .6 : lko.9 16 6 .2 32.07 15 . 0 : 7 0 0 .8 858.8: 105.3 70.9 17 2 0 .1 : 33.61 ok-3 8 6 8 .0 6 7 8 . 6 : 17k .9 9 -9 18 9.9 37.17 99.3 16.5: 7 8 3 . 4. 1007.3: 129 .6 1*0 . ^0 19 1 6 . 6 : 1 0 .6 8 0 6 .0 1208.9: 133.5 127.7 20 1*0.83 92.9 19»3l_ 1 2 .3 5i6.9 755.6: 99.7 21 13.0 1 * 9 .3 7 97.8 .25.3i_ 6 0 2 .8 75l.St~ 1 5 2 .6 22 59.20 20.9: 1^*7 6 k6 .ii 6 9 2 . 2 : 135.0 139.8 62.67 23 .... . 2 3 . 2 : 1 7 .? 575.8 9 6 8 . 0 : 133.8 173.5 21* 6 3 .2 0 17.8 22 . 1 * : U1 0 .3 610.7: 91.9 1 0 8 .6 65.00 25.7: 2 0 .1 25 3 h 9 .6 703.8: 89.6 11*1.1 26 65 ,2 0 21.5 23. 0 : 5 3 5 .2 621.3: 1 2 6 . eT 133.5 2 7 25.0: 25.1 69.53 U87.6 596.9: 121.7 1 5 0 .1 28 68.73 27.^: 2 L*.l I1I16 *5 720.1: 1 2 2 .8 173.7 29 69,83 23.9 9 6 .8 22.9: )in7.7 Ji05.lt 107.0 72.00 30 , 2 8 . 0 : 2 3 .3 356.9: 5 3 3 .6 : ioo.O 135.2 F in a l W eigh t P ercent Dry M atter TABLE 6 FIELD GARBOKI IRATE ROOT RESERVE STUDY ON FRUITING BROADLEAVED CATTAIL PLANTS DEVELOPING FROM GROWING S IT E INUN­ DATED BY WATER Lake : Height Surface: above Number Ho0 HoO : Temp *Surface : °F : cm 3-20-50:- Li.5.8 : 10.2 1 2 3-27-50: 51.8 : 12.7 k-3-5o : 53.1 : lh.O 3 L— 10— 50: 5k. 2 : 27.3 _ a 1.-17-50: 52.2 : . 3.2.7.. ....... 5....._ 6 W k-50: 57.5 : 56.9 7* 5-1-50. : 51l..2 : 58.0 8 5-8-50 : 52.5 91.0 5-15-50: 6 6 . 0 : I 0 6 .O 9 10 5-22-50: 7 6 . 5 : 137.2 5-29-50: 6 6 . 1 : 122.3 llb 12 c 6-5-50 : 7k. 0 : 130.7 6-12— 50: 7 3 .0 : lk5.9 lkd 6-19-50: 75.0 : l k l . l 6 - 26 - 5 0 : 8 k. 0 : 138.U 15®... 16* 7-3-50 : 8U.5 : lk 0 . 8 7— 10— 50: 80.5 : 155.1 17 . 18E 7-17-50: 71.5 : 165.9 7-2U-50: 70.0 : 1L8.7 19 20 : 7-31-50: 72.5 : 15k. 0 21 8-7-50 : 78.5 : I62_.k_ 22 8-lk-50: 79.5 : 166.U 8-21-50: 66.3 : 170. 8 23 8— 28 — 50: 53.0 . .: 167.9_ 2k 9-5-50 : 76.5 : 171.L _ 25. 9-11— 50: 57.0 : 159.6 26 9-18-50: 6 1 . 6 : 155.7 27h 28 9-25-50: 60.5 : l 6 k.k 10-2-50 : 55.0 : 166.9 29 10-9-50 : 5k. 0 : 155.2 ... Specimen Date Final Length cm Final Width cm Seed Female:Male Female Leaf Stalk Spike : Spike Spike Leaf 2 1 .1 1 .0 3 2 .2 1 .2 19.3 39.1 _35^9_ 80.7 7k.2 1 1 3 .0 120.5 l-g l t l .6 l.£ 158.9 17k.1 1.7 l6 l.7 131.6 lk.7 : 13.6 0.9_ 1-5 1 6 0 .1 lkk .2 . 2 -k : 1 7 .8 1 . 0 1.5 1 .6 1 .6 176.iL 16 k .k 17.1 : 1 6 .2 i.6> 166 i 5 l5k.9 Ik.k : 17oO 2 . 0 1 .6 175.2 1 . 8 1 7 9 .2 11.5 : 11.3 13.8 : 1 1 .2 2 .2 1 .5 186.5 1 7 8 .1 2 .6 1 5 8 .6 153.1 12.7 : l.k l.k 173.1 170.1 12.5. : .9.7 2 .8 191.8 185 .k lk.O : 1 0 .8 2 .8 l.g 2 .6 1 9 8 .8 189.7 13.5. : 1 .6 2.7 197*9. 188.3 12.7 : 193.7 182 .5 12.5 : 2.7 1 .8 186 .6 2 .8 1 .6 13.7 : 205.9 1 9 6 .8 2 . 8 lk .6 : 1 .6 213 .k 2 0 1 .6 19k.2 13.9 : 1 .6 __ 2 .8 2 1 0 .2 202.7 l k .2 : 1 .6 2.7 207.2 1 9 8 .6 lk.3 : 2 .7 l.k 2 0 1 .6 196.1 l k . l : 2 .6 1 .T1 C attail tops frozen back 6 to 8 inches» Green c a tta il leaves knocked down to water surface by snow, Broad-leaved c a tta il fru itin g bodies beginning to appear, Broad-leaved c a tta il pollinating heavily, Broad-leaved c a tta il pollination 90 percent completed, Broad-leaved c a tta il pollination completed. Female spike black brown . g Broad-leaved c a tta il vegetative tip s browned. a b c d e f : Percent : Carbo— : hydrate Specimen OvenFresh g Number Oven-Dry R dry Shoots Roots Shoots Roots Shoots Roots: Roots 3606 1 1 3 1 .7 2 3 0 . 7 8 .5 _6_.5_ 1 6 .7 : 5 7 .0 3 5 1 .7 0 6 I 4 ..O 1 5 . 0 : 2 U25.6 ^>.9 2 3 .7 3U5.3 9 . 8 U3.8 . 1 5 .2 : . 5 2 . 1 3 1 .9 3 ... 19 . h 2 8 7 . 3 6 9 .8 7 .2 U 1 0 .7 15 «U: U8.U0 lU 9 .li U53.3 1U.0: 5 0 .7 0 3 1 6 .8 UU.U 5 ... - " W .9 6.'5 _ 7 .5 6 U 6 . 6 0 7 . 6 5 2 .8 3 2 .6 1 3 .1 : U27.3 U02.7 1 5 .2 : 1+6 . h i 7 6 .9 7 1 3 . U l l 6 6 .3 U9.3 1 7 7 . I 8 5 5 .8 1 0 .5 : 7 9 0 .0 5 2 8 .3 7.1 5 5 .3 h-0.9 3 7 .8 8 2 .I T 9 7 . 0 1 0 .7 : 1C6U.2 9 1 1 .0 U0.37 9 10 1 0 .0 : 5 7 -0 32 .UO 1 0 .5 915 .u 5 6 6 0 8 96 .3 11 2U.80 69 . 6 1 1 1 .6 1 1 2 6 .it. 835 .9 8 .3 : . 9*9 12 8 .8 : 9U.2 6 5 .0 1 1 .3 .. 25 .1 3 8 3 3 .U 7 3 8 .0 2 0 .5 0 3 0 .6 1 6 .0 13 9 .9 : 9 5 9 .5 3 0 9 .7 15U .1 Percent Dry Matter Final Weight _______ . ___________________ 1 2 2 9 .3 5U5.8 108U.5 7 9 0 .3 1 0 0 0 .2 T 7 5 ^ “ 9 7 3 .3 2 7 7 .6 123U.3 UUo.2 9 3 5 .9 J4.6 0 . 6 1 0 8 3 .9 6 0 0 . 3 7 3 2 .7 3 6 6 .7 7 l 5 . 7 3U2.0 376.£ 1 0 U8 .6 685 .1 3 2 0 .5 7 5 5 .9 U36.3 8 5 7 .2 “ 3 6 5 :? 7 5 6 .1 f e l j f 6 9 6 .9 5 9 k . 1 7 7 6 .1 5 0 9 . 5 1 3 3 .1 1 7 7 .2 1 3 8 .5 1 8 3 .6 23U.9 1 9 2 .7 2 ^ 9 .3 20U.1 1 5 2 .9 2 7 3 .6 1 7 8 .2 189 .It 2 1 8 .3 2 0 7 .2 2 1 8 .2 2U6.6 " 767.2 h 39 2.9 ' W O T U i .o 75 .U 3 6 .5 2 7 .3 Uo.5 5 3 .3 2 2 .3 U5.9 3 2 .5 5U.3 4.2 •£ 7U.6 U6.7 7 9 .8 9 6 .0 103.5" 6 7 .9 1 0 .8 _ 7.5_: 1 6.3 . 9 .5 :. 1 3 .8 7_.7_: 9 .8 : 1 8 .8 3 1 .0 . 9 • 2 1 1 .6 : 2 0 .6 12 .0 : 2 3 .0 1 1 .9 : 2 7 .9 . 9 .6: 21 .U 26.1 1U.U: 2 6 .0 1 3 . U: 2 5 .0 1 7 .1 : 1 3 .0 : 2 5 .5 27. U 1 7 .3 : 1 6 .2 : 3 1 .2 3 1 .8 2 0 .3 : 1 7 .2 : 3 3 .7 Considerable yellowing in c a t t a i l s . 2 1 .2 7 2 7 .0 3 21+.90 30.1+0 33.50 3 1 .0 3 30 .So 33.53 3 2 .1 7 I16.IO 1? .30 s i.k z U .07 51.52. 6 1 .0 3 36 . uo 1U 15 16 17 18 19 20 21 22 23 2? 26 27 28 29 30 • ------. - .TABLE 7 FIELD CARBOHYDRATE ROOT RESERVE STUDY ON MIXTURE OF FRUITING AND NONFRUITING BROAD-LEAVED CATTAIL PLANTS DEVELOPING FROM GROWING SITE NOT INUNDATED BY WATER Specimen Number 1 2 3 5 5______ 6 7a 8 . 9: 10 iib 12G 13 i5d 'l5>e , ... 17 lte 19 20 21 22 23 25 25 26 27h 28 29 OP________ Date 3-20-50 3-27-50 5-3-50 L-io-50 5-17-50 5-25-50 5-1-5P. 5-8-50 5-i 5-5Q 5 - 22 -5 0 5-29-50 6- i - ^ 0 6-12-50 6-19-50 6 - 26-50 7-3-50 7 - 1 0 -5 0 7-17-56 7-25-50 7-31-50 8-7-50 8-15-50 8-21-50 8-28-50 9-5-50. 9-11-50 9-18-50 9-25-50 10-2-50 10-9-50 Lake Height Surface above Ground H2° Temp Surface cm °F 55.8 51.8 53.1 55.2 . 3.3 . 5 2 .2 37.7 15.7 57-5 55.2 25.9 52.9 52.5 6 6 .0 Sk.8 . 76.5 66*1 (-1-9 6 7 .0 75.0 73.0 8 ^ .8 75.0 115.3 85.0 1 1 5 .6 82.5 1 1 9 .0 80.5 157.8 71.5 139.9 70.0 135.1 72.5 170.1 78.5 157-7. 79.5 167.2 66,3__ l & J . 53.0. 162.3 76.5 153..2 . 57.0 162.6 6 1 .6 151.8 66.5 153.3 55.0 155.9 5U.o 155.8 Final Length cm ;Final Width : cm » « ?Seed Female Male :Female Leaf:Stalk Spike Spike: Spike Leaf • • •• • • •• • •• 15.1 : . 25-7:________ 51.5: 5 6 .6 : 65.7: 86 *0 : 69.2: 108.7: 1 2 5 .2 : 97.8: 1 3 0 .1 : 153.5: _ _ J_ 15.1 133.2:156.5 151.2:156.6 r 11 , 5. 165.2: l55.9 1 1 .9 150.8:151.7 1 2 .0 158.8:159.6 1 1 .9 162.5:156.5 1 1 .3 163.1:156.0 1 2 . 2. 156.6:152.3 1 5 .8 150.5:139.6 1 1 .1 151.8:150.8 1 0 .7 136.6:155.7 1 1 .7 138.2:135.8 1 3 .1 122.6:155.9 1 1 .8 •• • • • • • » • • • •» « • • •• • « 12.8 : 11.5: 10.2: 16.5: 7.9: : : : : : .... . • : : : 1 .0 1-3 1.5 1*5 1 .1 1.5 1 .5 2.2 2.8 2.7 2.9 2.8 3.0 2.9 2.8 2 .2.. 3.0 2.9 2.7 1 .3 1 .5 1 .5 i-3 1 .5 i-5 1 .5 i.5 1 .5 1 .3 1 .2 1 .2 a Cattail tops frozen Lack 6 to 8 inches. b Green c a ttail leaves knocked down to water surface by snow, c Broad-leavedfruiting bodies beginning to appear, d Broad-leavedc attail pollinating heavily, e Broad-leavedcattail pollination 90 percent completed, f Broad-leavedcattail pollination completed. Female spike black brown. Final Weight Fresh g Shoots Oven-Dry g Roots Shoots Roots Shoots 200.6 . 5.2,3 .215 , . 27.3 29.3 llS.7 5 0 0 .1 2 0 6 .k U19.8 350.7 3 6 0 .0 1106 .3 550.0 6 k6 .6 Os sQ . rn -d 753.2 6 b2 .3 7&U.9 623.1 6 3 6 .O 6 1 1 .8 593.6 " W .S ’ 521.3 kk5 .k 5 0 7 .5 362 .ii 2i4b.lt g h Roots:Shoots Roots:Shoots 2 0 0 .6 : 52.3: 215.6: 50.7: 117.8: 3U.U: 99.6: 27.7: 2 U.0 : 9.9 90.0: 2.7 215.6: 3 .2 U3.lt 10.9 8 2 .U: 8 .6 399.0: 10.0 U7.0: 7.2 38k.0 : 35).9 30.2: 7.9 225.7: 1 6 .2 29.1: 1 2 .0 217.5: 50.5 3 7 . 2 : 1 1 .1 305.1: 53.IT 87.3: l 6 . 0 U07.9: 56.2 56.8: 22.3 278.5: 8 0 .2 6 8 . 0 : 1 U.U 566.5:1^9.3 72.3: 19.9 Jj.25. 8 :109 .9 53.6: 17.6 kOO.k: 113.9 59.6: 22.3 U22.9:211.3 102.7: 20.1 6 0 I1.i 1.: 1 5 1 .1 7U.8: 23.6 k05.5:l5k.l 99.9: 2 U.6 5 0 2 . hi 1 9 2 . a k9 2 . 8 : 1 6 6 .2 103.1: 26.7 123.0: 26.9 528.2:170.9 L.58.6:153.U 10U.9: 25.9 h i ? . 3 =17 b-: n ~ 113.7: 29.7 80.5: 3 U.2 286.2:169.7 117.2: 30.U k08 . 2 :158.3 h39.7:lii.6. 6 13U.5: 33.2. 111.3: 32.3 Li05. 6 :163.7 98.5: 3 U.0 395.5:123.k 9k.9: kl.2 327.6: 99.1 ercent Carbolydrate Ovendry Roots Roots 2 6 .1 23.-5. 2 9 .2 27.8 26.7 2 0 .0 20.7 1 2 .2 13. k 13-k 1 2 .2 2 1 .it 2 0 . b1 2 .0 1 7 .0 13. k lk .1 1 7 .0 18. h 19.9 2 0 .2 23.3 22.9 27.1 2 8 .1 28.7 30.6 27.k 2k.9 29.0 Specimen Number 55.17. 5k. 00 U7.37 58.13 U3.80 U3.31 k9 .3 0 37.87 kl.77 25.93 27.23 36.93 U5.90 3U.57 kO.60 38.80 U7-13. U8.03 k6.80 k9.6 7 50.53 60.17 59.27. 61.57. 59.17 60.87 62.33 6h.3_0 5 7 -6 I_ 59.53 Broad-leaved cattail vegetative tips browned, Considerable yellowing in cattails. 1 2 3 k 5 ............................ 6 7 8 9 10 11 12 13 lk 15 16 17 18 19 20 21 22 23. ...... 2k 26 27 28 29 30 . ... March 20, 1950, and continued through October 9, 19^0. For the Denver area, minimum carbohydrate root reserves occurred, in general, from May 22, 1950, u n til July 31, 1950, for the broadleaved c a tta ils developing from the growing s ite inundated by -water. For the narrow— leaved c a tta ils developing from the growing s ite inundated by water, the period between June 5, 195>0, and July 2 l+, 1 9 5 0 , was one of low root reserves. The period May 8 , 195’0, to July 3, 1950, was one of low root reserves for the broad-leaved c a tta ils developing from the growing s ite not inundated by water while the period July 3, 1950, to July 2U, 1 9 5 0 , showed low root reserves for the narrow-leaved c a tta ils developing from the growing s ite not inundated by water. I t is well known th at growing seasons vary, not only according to altitude and la titu d e , but from year to year as well. Accordingly, while these dates in general lim it the 1950 periods of low root reserve for the c a tta ils developing in the Denver area growth situations they could not be used to determine low root reserves for c a tta ils develop­ ing in other sections of the western United States. Notes Pertaining to Appearance of Fruiting Bodies, Pollination Time, e tc . Study of c a tta il in th is area showed that when plants develop from a growing s ite inundated by water fruiting bodies do occur. I t should be mentioned here that the broad— leaved c a tta il s ite not inundated by v;ater had a drainage ditch passing through i t which carried water to a depth of six inches immediately following the occasional 12 heavy spring rains in the area. In no instance, however, did water remain in the drainage channel more than 2 U hours a f te r a downpour. No fruiting bodies developed and, accordingly, no pollination occurred in the narrow-leaved cattails developed from the growing s ite not inundated by water. Weekly Water Temperatures Approximately Eight Inches Below Lake Surface I t was interesting to note that c a tta il growth in the water inun­ dated growing sites began when the water temperature reached [|5>.8° F. In the developing sites not inundated by water growth development came la te r . This growth lag appeared to be associated with lower s o i l temperatures. was appr The narrow-leaved cattail s i t e not inundated by water imately eight weeks behind the water inundated one in plant development. After each snow and rain storm there was a noticeable drop in lake water temperature. Nevertheless weekly temperatures increased steadily up to the la t t e r part of June when a temper*ature of 81*.0° F was recorded. Warm temperatures held relatively steady u n t i l the middle of August when there was a noticeable gradual decline to the f in a l recording of ^U.0o F on October 9 , 195>0. Height of Plants above Water Surface or Ground Line Plants developed from the grov/ing s ite s inundated by water were at maturity approximately 30 - 5*0 cm taller than th e ir counterparts developing in non— inundated growth sites• 13 Nonfruiting broad-leaved plants developing in inundated growth sites were approximately lj.0 cm t a l l e r than fruiting plants developing in similar s ite s . In the narrow-leaved and broad-leaved c a tta il growth s ite not inundated by water root reserves were a t a minimum when the plants had attained a height of approximately 1 0 0 -1 3 0 cm and 5 0 -1 2 0 cm above the ground line respectively. Total Length of Plant Shoots At maturity, leaf lengths were approximately 30 cm more for non­ fruiting than for fru itin g broad-leaved c a tta il plants. Leaves on plants developed from the growing sites inundated by water were approximately 5 0 - 7 0 cm longer than on similar plants develop­ ing in non-inundated growth s i t e s . Total Length of Seed Stalk At maturity, in the growing site s inundated by water, leaf length extended beyond the seed stalk approximately 10 cm in the broad-leaved c a tta ils and 3 0 -U0 cm in the narrow-leaved c a tta ils . In general, for both c a tta il species the period of seed stalk elongation was one of low carbohydrate root reserves. Length of the Female and Male Spikes Female and male spike lengths did not increase noticeably from time of th eir i n i t i a l field appearance u n til they had attained f u ll maturity• Ik Width of Female Spike The broad-leaved c a tta il female spike increased rapidly in diameter from an in itia l 0 .9 cm to the maximum diameter of about 2 .7 cm, an increase of about two cm over an appro:cimate seven-week period. The narrow-leaved c a tta il female spike likewise increased rapidly in diameter from an in itia l 0 .5 cm to the maximum diameter of about 1 .7 cm, an increase of about 1 .2 cm over an approximate six-week period. Both species of cattails go through a pronounced depletion of carbohydrate root reserves during this period of rapid expansion of the female spike. Width of Leaf Broad-leaved c a ttail leaves increased rapidly in width from an in itia l 1 .0 cm to the maximum width of about 1<>5 cm, an increase of about 0 .5 cm over an approximate eight-week period. Narrow-leaved cattail leaves likewise increased rapidly in width from an in itia l 0 .5 cm to the maximum width of about 0 .8 cm, an increase of about 0 .3 cm over an approximate six-week period. I t was interesting to note that, in general, both species of cattails attained maximum leaf width immediately preceding their respective periods of low carbohydrate root reserve. Shoot and Root Fresh Weight The shoot and root fresh weight samples were limited in size to the amount of cut plant material which could be placed in a six-inch cubical wire basket. Percentage Dry Matter in Shoots and Roots In a ll five c a tta il growth situations investigated there was a marked tendency for percentage dry matter in roots to be relatively high when percentage carbohydrate in roots was high and similarly to be low when percentage carbohydrate was low. In contrast to the roots, the percentage dry matter in the shoots, in the five c a tta il situations investigated, increased apparently at the expense of the carbohydrate root reserves which a t the same time were being depleted. I t is apparent that root reserves are at a minimum for the fruiting and nonfruiting broad-leaved c a tta il plants developing in growing sites inundated by water when the percentage dry matter in the roots is approximately 10 percent. Likewise, root reserves are at a minimum for broad-leaved c a tta il plants developing in growing sites not inundated by water, and for the narrow-leaved c a tta il plants developing in both the water inundated and non-inundated site s, when the percentage dry matter in the roots is approximately 1 £ percent. I t can be seen that by making laboratory fresh, and oven-dry, weights on c a tta il root samples and calculating percentage dry matter in the same, i t would be possible to follow the percentage carbohydrate in roots through the growing season without performing the more timeconsuming, but more precise, chemical analyses. Percentage Readily Hydrolizable Carbohydrate in Oven-Dry Roots Pitying the winter dormancy period c a tta il roots had a carbohydrate 16 root reserve of approximately 60-70 percent. As plant development progressed in the three broad— leaved c a tta il growth situations, a minimum root reserve of about 25> percent was reached. This minimum was followed by a gradual increase of food reserves which continued to the end of the growing season. In the two narrow-leaved c a tta il growth situations a similar de­ crease in root reserves occurred as plant development progressed, fo l­ lowed by the gradual increase of reserve food which continued to the end of the growing season. I t was interesting to note that the minimum readily hydrolizable carbohydrate root reserve obtained for the narrow-leaved c a tta il was about UO percent. Field observations indicated th at the narrow-leaved c a tta il roots were more laden with starch than were the roots of their frequent fie ld associates, broad-leaved c a tta il. This work indicates th at maximum c a tta il eradication and/or control with minimum time, effo rt, and cost should be effected i f the herbicidal applications are made: (l) between the time of f i r s t appearance of the fruiting stalks and when pollination has been com­ pleted, ( 2 ) when narrow-leaved c a tta il plants developing in growing s ite not inundated by water attain a height of 100—130 cm, (3 ) when broad-leaved c a tta il plants developing in growing s ite not inundated by water attain a height of £0 - 1 2 0 cm, (U) during the period between appearance and maximum seedstock elongation, ( 5 ) between i n i t i a l appearance and when broad-leaved c a tta il female fruiting body attains a diameter of 2 .7 cm, ( 6 ) between i n i t i a l appearance and when narrow- 17 leaved c a .tta il female f r u itin g body a tta in s a diameter of 1 . 7 ( 7 ) when percentage dry matter i s cm, approximately 1 0 percent fo r roots of broad-leaved c a t t a i l developing in s i t e s inundated by w ater, ( 8 ) when percentage dry matter i s approximately If? percent for roots of c a t t a i l s developing in the growth categories other than in 7 above, ( 9 ) when a minimum carbohydrate ro o t reserve of percent i s reached for broad-leaved c a t t a i l p la n ts , reg ardless of the growth s itu a tio n ( 1 0 ) when a minimum carbohydrate roo t reserve of percent i s reached for narrow— leaved c a t t a i l p la n ts , regardless of the growth s itu a tio n • 18 IDENTIFICATION OF SPECIES Differences in effect of herbicides on -water plants can be accounted for, in part at le a st, by variation in species or varieties of species present at the time of treatment. Translocation studies were performed on the following species and varieties of plants (Coulter and Nelson U, Fassett 8, Fernald 9, Femald 1 0 , Hotchkiss and Dozier 15>, Moore 2 2 , Muenscher 23, Ogden 2h> Robinson and Fernald 26, Rydberg 27, St. John 2 8 , Schmeil and Fitschen 30) which personal observations have shown to bo some of the most troublesome vascular plant waterweeds which impede irrigation water flow in the United States west of the 100th meridian: water sedge, Carex aquatilis Wahl. leafy pondweed, Potamogeton foliosus Raf. var. genuinus American pondweed, Potamogeton nodosus Poiret gigantic sago pondweed, Potamogeton pectinatus L. slender sago pondweed, Potamogeton pectinatus L. Richardson's pondweed, Potamogeton richardsonii (Ar. Benn.) Rydb. narrow-1eaved c a tta il, Typha angustifolia L„ broad— leaved c a tta il, Typha la tif o lia L. horned pondweed, Zannichellia palustris L« I t w ill be noted that distinction has been made between slender and gigantic sago pondweeds, both taxonomically known as Potamogeton pectinatus L. 19 I t has been observed in the laboratory that the coarse form, so common in flowing waters of the west, is much more re sista n t to chemicals than is the slender form, more common in the relativ ely quiet waters of the west. Thus, i t seems desirable in th is study to make a d istin ction between the two forms of Potamogeton pectinatus L. 20 MATERIALS AND METHODS The methods and materials used in making herbicidal treatments can in this study, for convenience, he divided into five sections. I. Leaf— Tip Immersion Test In the le a f-tip immersion te s t (see plates I and I I ) , one cut leaf tip from one plant in each beaker was immersed in commercial undiluted phenoxyacetic acid derivative for 2U hours. The immersed part was then cut off with scissors and the plant placed in the green­ house for a five-week period. At the end of this period, the amount of living and dead leaf material in immersed leaves, non-immersed leaves on the same plant, and leaves developed from la te ra l buds, were measured separately in each beaker for length and the combined living and dead material used for weight determinations. Killing effect on the leaf of which the cut tip was immersed, or on the non— immersed leaves on the same plant, was considered to be of a local type. Killing effect on the leaves developed from la te ra l buds after the immersion treatments, and absence of new re­ growth in the eight— week period following leaf harvest was considered to be evidence of a permanent or systemic type of k i l l . Plants used were grown in beakers from la te ra l buds and were approximately 2 - 1 / 2 feet t a l l a t time of te s t. The immersion solution Is liste d for each group of five beakers. The f i r s t two numbers in each group of five, e.g ., 1— 2, 6— 7, 11 — 12, 21 e tc ., are treatments in which narrow-leaved c a t t a i l plants were used. The l a s t three numbers in each group of fiv e , e .g ., 3— 5, 8 - 1 0 , 1 3 - 1 5 > e tc ., are treatments in which broad-leaved c a t t a i l plants were used. The arithm etical average for each group of plants treated , i . e . , narrow-leaved c a t t a i l and broad-leaved c a t t a i l , is reported in such a manner th a t the percentage k i l l expressed in length and weight can be compared with the non-treated control group which are numbered 35-UO. II. Root— Immersion Test In the root-immersion t e s t (see p late I I I ) , roots of the plants were immersed in a solution containing 10 ppm of phenoxyacetic acid compound for 2b hours, rinsed in three changes of tap water and the plants then put in beakers containing tap water, which in turn were placed in the greenhouse for a five— week period. At the end of th is period, the amounts of living and dead leaf material in each beaker were measured for length and weight. Killing effect on the leaves of these treated plants was considered to be of a local type. Killing effect, as evidenced by absence of new re - growth in the eight— week period following le a f harvest, was considered to be evidence of a permanent or systemic type of k i l l . As in the previous t e s t , plants used were grown in beakers from la te r a l buds and were approximately 2 — 1 / 2 fe e t t a l l a t time of te s t . As in the le a f— tip immersion t e s t , the immersion solution is lis te d for each group of five beakers. 22 The f i r s t two numbers in each group of fiv e , e .g ., 5> 1— 5>2 , 6 1 — 6 2 , e tc ., are treatments in which narrow-leaved c a t t a i l plants were used. The l a s t three numbers in each group of fiv e , e .g ., 5 3 - 5 5 * 63-65, e tc ., are treatments in which broad— leaved c a t t a i l plants were used. The arithm etical average for each group of plants treated , i . e . , narrow— leaved c a t t a i l and broad— leaved c a t t a i l , is reported in such a manner th a t the percent k i l l in length and weight can be compared with the non-treated control group which are numbered 8 1 - 8 5 . III. Aerial Herbicidal Treatment on Plant Materials Transplanted to Buckets of 5-Gallon Capacity I t was desirable to know something about the regenerative powers of broad— leaved c a t t a i l p lan ts. In order to investigate th is question, 15 basally clipped stocks were measured for bud lengths and individ­ ually placed in 3 ,0 0 0 nil beakers, containing only tap water, on October 2 6 , 19U9. The f i r s t crop of new growth from the buds was measured, recorded, and harvested December 3 0 , 19U9 • °n th is and successive dates the number of new buds was counted and amount of growth developing from the new buds was measured, recorded, and harvested. The length of the longest leaf in each new plant developed from the old stock was recorded as shoot length. On October 18, 195°* bbe rhizomes were processed, as discussed e a rlie r in the section on seasonal carbohydrate root reserve trends in roots and rhizomes of broad—and narrow— leaved c a tta ils , and the percentage hydrolizable carbohydrate determined as described by 23 Woodman (37) and Schaefer and Hartmann (31). In this determination for reducing sugars, i . e . , reducing substances, the monosaccharides, the disaccharides lactose and maltose and upon hydrolysis sugars yielded by starch, pentosans and various hemicelluloses possess the property of reducing alkaline solutions of metallic s a lts , such as copper or mercury, oxygen being •withdrawn and the metal precipitated either as such or as a lower oxide. Chemical analyses thus showed the amount of readily available carbohydrates s t i l l present in c a tta il rhizomes after between 1 S > and 20 new and complete plant en tities had emerged from the several rootstocks. Before making herbicidal applications on aerial parts of field grown, broad- and narrow-leaved c a tta ils , the plant materials were transplanted from the fie ld directly to buckets of five-gallon capacity one week before treatment (see plates VI and VII), and with as l i t t l e root disturbance as possible. The inside of the containers was painted before use with two coats of coal-tar paint to in h ib it ru st formation. Exposed surface area measurements were calculated to be 79 square inches for each of the five-gallon buckets. The buckets containing the transplanted material were f ille d with tap water and the shoot lengths above the surface of the water in each bucket were measured. In itia lly , percentage dead plant material, based on length and weight measurements, was considered a measure of the temporary lethal effects. Absence of green shoot development after treatment and a 2k relatively low percentage carbohydrate, i . e . , less than 30 for broad­ leaved c a tta il and less than U0 for narrow-leaved c a tta il, root reserve three or four weeks after treatment were considered indica­ tive of lasting leth al effects for any particular herbicidal treatment. At the termination of th is particular study i t became apparent that in a number of instances treated plants with no green shoot regrowth development three and four weeks after treatment frequently had a relatively high percentage carbohydrate root reserve, and, therefore a considerable regrowth potential. Aerial herbicidal application was made in each case with the one-quart capacity, model A, Sure Shot pneumatic sprayer. The quantity of the particular chemical used in this study was determined by computing the difference between the i n i t i a l amount placed in the sprayer and the amount remaining therein after aerial herbicidal spray application had been made. I n itia l a ir pressure in the sprayer for each spraying was 120 pounds per square inch. Bohmont (2), Grigsby, Churchill, Hamner and Carlson (13) have pointed out the fact that herbicidal a.ctions of 2 ,U— D and 2 ,U— 5 -T are related to the actual acid equivalent of the formulation. Accordingly, systemic herbicidal dosage amounts are, in this study, stated on the acid— equivalent basis. The chemical compounds used in the various aerial herbicidal sprays were formulated as follows: 1. Alpha benzene hexachloride—300.0 g made up to 2,000 ml with Xylene containing one percent California Spray Chemical Ortho emulsifier no. 5 2^ 2. Amine s a lt of 2,U-D~2 *U9 g made up to 1 ,0 0 0 ml -with ■water 3. Anhydrous copper sulphate—106.6 g made up to 2,500 ml ■with water h. Ester of 2 ,U-D—2 .0 8 g made up to 1,000 ml with water 5. Ester of 2, U, 5-*T— *2 .0 8 g made up to 1,000 ml with water 6. Pentachloropheno1 —2 0 0 .0 g made up to 3 ,0 0 0 ml with xylene containing one percent California Spray Chemical Ortho Emulsifier no. 5 7. Polyethanol Rosin Amine 0500 (D~U) 255 ml Rosin Amine D 1;5 ml Rosin Amine D acetate 229.9 g Made up to 2,500 ml with xylene 8. Polyethanol Rosin Amine 1100 (D-12 255 ml Rosin Amine D ml Rosin Amine D acetate 239.2 g Made up to 2,500 ml with water 9. Sodium arsenite—185.0 g made up to 2,500 ml with water . 11. Sodium chlorate—750oO g made up to 2,500 ml v/ith water 10 Sodium pentachlorophenate—250.0 g made up to 2 ,0 0 0 ml with water . Sodium s a lt of 2,U-D—U.ll g made up to Lj.,100 ml with water 13. Sodium s a lt of 2,U— D— -8.23 g made up to U.,100 ml with water 1 U. Sodium s a lt of 2,U-P—*l6.1|6 g made up to U,100 ml with water 15. Sodium sa lt of 2,U-&--3 6 .8 6 g made up to 3*785 ml with water 16. Sodium trichloroacetate—2 7 9 .8 1 g made up to 3*785 ml with water 17. Xylene—1,980 ml to which was added 20 ml (making one percent solution by volume) California Spray Chemical Ortho Emulsifier no. 5* making a to ta l volume of 2 ,0 0 0 ml 12 26 W e ttin g a g e n ts u s e d w e re a s f o llo w s : 1. Span 20 and Tween 20 (products of Atlas Powder Company) 2. Span 85 and Tween 85 (products of Atlas Powder Company) 3* Vatsol 0T— B (product of American Cyanamid Company) Immediately a fte r the aeria l spray treatments, the water in the buckets was siphoned out of the containers with l / 2 -inch rubber tubing, the buckets r e - f ille d with water, siphoned out a second time, and again r e - f ille d with water. Flooding of the buckets continued for one minute after they were f u l l to wash out any chemical th at might have fallen into the bucket during spraying. Thus, any effect on the root system of the c a tta ils would be due to translocation effects from the above­ ground or above-water plant p arts. After the a e ria l herbicidal treatment had been made on the broadand narrow— leaved c a tta il plants they were allowed to remain in their containers on a concrete loading platform located on the north side of a building, in one experiment for three weeks, in the other for four weeks. At the end of th is period of time the amount of living and dead leaf material in each bucket was measured for length and weight. Consideration of the effects of temperature differences on plants in the two te s ts made i t desirable to disregard the number of green shoots developing a fte r treatment as a measure of permanent leth al effect on the treated plants. Percentage hydrolizable carbohydrate root reserve three and four weeks after treatment appeared to be a re lia b le manifestation of permanent injury brought about in the plants 27 receiving the various herbicidal applications. Measured percent survival at the termination of these two tests •was determined by taking the arithmetical average of percentage dead based on to ta l length and to ta l weight measurements and then subtracting from 1 0 0 . These figures, as suggested e a rlie r, were con­ sidered a measure only of temporary effects on the treated plants. IV. Aerial Herbicidal Treatment on Plant Materials Transplanted to Metallic Containers of 27-1/2 Gallon Capacity In the aerial herbicidal treatment on field grown plant materials transx^lanted a week before treatment into metallic containers of 27-1/2 gailon capacity (see plates VIII, and XII-XXVII), most of the true waterweed, American pondweed, horned pondweed, leafy pondweed, Richardson1s pondweed, gigantic sago pondweed, and slender sago pond­ weed plants used were transplanted from the fie ld directly to the container with as l i t t l e root disturbance as possible. In some in­ stances tubers of gigantic sago pondweed were planted d irectly into the so il at the bottom of the tank. Approximately ninety 27— 1/2 gallon and six 7— 1/2 gallon metallic containers were used in this study. The larger capacity containers were prepared by cutting 5 5 — gallon metallic drums in half crosswise or lengthwise. The smaller capacity containers were prepared by cutting 15-gallon metallic containers lengthwise. The inside of the containers was painted before use with two coats of coal-tar paint to 28 in h ib it r u s t formation. The exposed surface area measurements were calculated to be 3 8 0 square inches for the large round tanks, and 7^9 square inches for the large long tanks. The small tanks (cut from the l^-gallon containers) viere calculated to have an exposed surface area of 336 square inches. These smaller tanks were used only for the untreated co ntrols. Before each of the various chemical treatments were made, water was siphoned out of the tanks with l/2 — inch rubber tubing. Because the s o il in the bottom of the tanks was not p erfectly le v e l, small water puddles, up to two inches in depth, remained in the bottom of the planted tanks. This condition, i t i s believed, simulates th a t which would p revail when water flow i s completely cut off in an ir r ig a tio n d itch , i . e . , complete drainage would not occur because of occasional low spots in the ditch bottom. A erial herbicidal applications on the plants in the drained tanks were made with the one-quart capacity model A, Sure Shot pneumatic sprayers. The spray was applied on the flaccid aquatic plants which, in the absence of the usual water supporting medium, were lying on the s o il in which they were rooted. The sprayed plants normally submerged were allowed to remain lying on the s o il a t the bottom of the tank for a period varying from two to six hours following spraying, to allow the herbicides to penetrate the plant tissu e s. In the report th is i s recorded as exposure time. At the end of th is period of time, the tanks with the sprayed 29 plants were f i l l e d with tap water and drained immediately. This operation was repeated twice a fte r which the tanks were f i l le d a th ird time with tap water and flooded for five minutes. The tanks then stood for seven days approximately one meter d ista n t below a lighted 7 ^0 — watt incandescent lamp. After the seven-day period under a r t i f i c i a l lig h tin g , the water in the tanks was siphoned off and the tanks and plants were moved to an outside sunny exposure next to the greenhouse. The tanks were again f i l l e d with tap water and the plants observed for effects of the chemical treatments. Water was drained off of the chemically untreated control tanks and the plants growing therein were subjected to the same two—to six— hour desiccation as the chemically sprayed tanks, the only difference being th a t no chemical was applied. I t should be noted th a t th is same uniform technique was used even when using contact chemicals, e .g ., copper sulfa.te, sodium arsen ite, etc . I t is believed th a t by f i l li n g and siphoning off the water in the tanks twice and then f i l l i n g and flooding for five minutes, any herbicide which had not entered the plant in the two—to six— hour exposure period would be washed out of the tank, thus, any effects observed after treatment could be a ttrib u te d to entry of the chemicals into the flaccid plants during the exposure time when the water was out of the tank. This washing effect was thought to simulate the wa.shing effect which would occur when water is turned back down an irrig a tio n ditch 30 •where rooted aquatic weeds have been sprayed, using a technique similar to the one described for the rooted aquatic weeds growing in the tanks. The amount of phenoxyacetic acid derivative used on these various tanks was calculated on an acid equivalent pound-per-acre and gallonper-acre basis. Contact herbicides used were calculated on a pound- per-acre and gallon-per-acre basis. In some instances, a contact herbicide was applied 2h hours after a systemic herbicide had been used. In other experiments, several weeks elapsed between the two or three separate treatment applications. In each case, one half of the plants in the control tanks was undisturbed and the remaining half of the plants in the control tanks was clipped to the ground with household scissors. Thus, a com­ parison can be made between no disturbance, clipping, and chemical treatment. At the termination of the experiment, the water was siphoned off and an estimation made as to the percentage of the so il area in the bottom of the tank supporting green plant growth of the i n i t i a l species present in the tank. The so il in the bottom of the tanks was examined for the presence of living underground propagules. From this physical examination estimated percentage survival is reported. ITIhen systemic herbicides were used on both f i r s t and second treatments, the summation of the two was reported as pounds per acre for that particular treatment. In like manner, when the same kind of contact herbicide was used 31 in both f i r s t and second treatments, the summation of the two was reported as pounds per acre for th at p articu lar treatment. In order to obtain a quantitative measurement of the effect of the various chemicals used, in several instances the living plant material remaining in the tank a t the termination of the experiment was harvested, weighed fresh, oven-dried, and weighed again. This quantity of oven— dried residue remaining was compared with the ovendried residue in the control tanks in which half the area was clipped each time an a e ria l herbicidal treatment was made on the experimental tanks. These data are presented with photographs of the tanks to which they pertain. V. Miscellaneous Tests I t was desirable to know whether 2 ,L|— D would move past the water lin e when emergent aquatic plants were treated with th is herbicidal compound. The following t e s t was then performed to determine i f 2 ,U— 1) could be detected in the underwa.ter roots of c a tta il plants, the leaves of which were sprayed with th is phenoxyacetic acid derivative. About 15> selected,pulverized c a tta il root specimens were ex­ tracted with benzene in a Soxhlet extraction apparatus. The benzene extract was evaporated to dryness in a t e s t tube; a few crystals of chromotropic acid ( 1 , 8 — dihydroxy naphthalene 3 >6 -disulfonic acid) introduced; two ml of concentrated sulfuric acid added, and the material heated in a glycerine bath a t 15> 0 ° C for 1-1/2 to two minutes. Freed ( ll) found th at when following th is procedure s. deep wine— purple color develops quite rapidly while heating when 2 ,U— D is present. 32 Water sedge plants -were treated (see plate XI) -with herbicidal concentrations of 2 ,U— D and 2,U*5>— T to observe the effect of these applications on root reserve depletion and leaf k i l l in the sedges. Accordingly,, a procedure -was followed similar to that used in trans­ planting and treating the c a tta ils except th at in th is instance two containers of 2 7 - 1 / 2 gallon capacity were used and, too, percentage k i l l was determined by weight measurements only. In the Appendix, photographs showing the various plant growths before and after systemic herbicidal treatment are presented together with photographs of mechanically dipped and untreated plants so that a visual evaluation may be made of the respective treatments. Photomicrographs of aquatic plant sections prepared (Corrington, 3, Sass, 29) by the paraffin and celloidin methods (see plates IX and X) are included to show variation in aquatic plant anatomy, particu­ la rly the marked difference in amount of cutin, number of stomata, location of chloroplasts, and degree of specialization of the vascular system. Certain anatomical features peculiar to the species (see plate IV) are shown which should be helpful in effecting accurate fie ld identi­ fications of the waterweeds. In this report a l l linear measurements, except where otherwise noted, are made in centimeters, weight measurements in grams, and temperature measurements in degrees Fahrenheit. 33 DISCUSSION OF RESULTS I. LeaT-Tip Immersion Test Results from immersing one cut le a f tip per c a t t a i l plant for 2k hours in commercial strength systemic herbicidal stock solution are presented in table 8 . These data show the changes in amount of living material in the above-ground plant parts brought about, five weeks after treatment, by the action of certain herbicides passing from the tip of the cut le a f through the plant body. The isopropyl esters of 2,U,5>-T and 2,U~D were more pronounced in th e ir le th a l effects on the narrow-leaved c a t t a i l than was the triethanolamine s a l t of 2 ,li— D. In the treated , broad-leaved c a t t a i l plants, on the other hand, the difference in le th a l effects between the ester and s a l t of 2 ,U-L was much le ss pronounced. Perhaps the most interestin g re su lts of th is t e s t l i e in the le th a l effects produced by the apparent passage of the esters into the leaves developed from the i n i t i a l l y undeveloped, basally located, la te r a l buds. I t appears th at the le th a l effect on these leaves was brought about by transmission of the herbicide through the immersed leaf tip past the waterline and into the crown of the plants involved. Associated with th is phenomenon is the fact th at no shoot regrowth developed in the ester treated plants in the eight-week period 3k TABLE 8 EFFECT OF IMMERSING ONE LEAF TIP PEE CATTAIL PLANT FOR 21+ HOURS IN COM M ERCIAL STRENGTH SISTEMIC HERBICIDAL SOLUTION DATA TAKEN £ W EEKS AFTER TREATM ENT Plants Treated and Compound Used Narrow— Leaved Cattail Typha anRustifolia L. No treatment Total Leaf LenRth Leaf with Immersed Nonimmersed Leaves _______TiP . ______ Beaker Dead Living Dead Living % Number cm Doad cm cm cm u~5 7 .0 6-7 1 39.1 0 .0 1 0 0 .0 2 2 9 .0 8 8 .9 2,1+-Dj Isopropyl ester 11-12 8 2 .0 0 . 0 1 0 0 . 0 1 6 I+.1 0 .0 2 ,L|-Dj Triethanolamine s a lt 16-17 Uo .6 9 1. h 1-3 0 .0 1 6 2 .2 8 -10 1 7 6 .3 2 U. 1 2,U*5>-T, Isopropyl ester Broad— Leaved Cattail Typha la tifo lia L. No treatment 2,l+*5-T, Isopropyl ester HI+.5 6 . 0 1 0 0 . 8 258.7 3 1 .1 1U5.3 0 .0 35.5 9 7 .9 516.5 87.5 315.5 U17.8 2 ,I|.-Dj Isopropyl ester 13-15 185.3 0 . 0 1 0 0 . 0 3U3.5 2 3 6 . 1 2,1+-D, Triethanolamine sa lt 18-20 188.8 0 . 0 1 0 0 . 0 282.5 3 1 1 . 8 Shoot Total Leaf Weight Regrowth Leaves Developed Dead % Dead 9-22-50 Living OvenOven- Oven- Maximum from Later a]L Buds Fresh dry Fresh dry dry Leaf i T ~ Dead Living G ram s G ram s Basis D ead cm Grams Grams Dead cm Length 0 .0 1 0 6 .0 0 .0 1 .1 O.U 9.0 2 .1 15.0 1 1 .1 7 0 .1 hh-2 193.3 2 3 .6 U.6 2 .U 6.5 1 .8 6o.5 0 .0 1 0 0 .0 8 2 .1 3b0.9 2 0 .0 3.U l.U 7.0 1.7 h h .3 0 .0 8 5 .8 0 .0 U63.5 0 .0 2 .0 0.9 1 1 .6 2.9 23.6 1 2 .0 12.3 0 .0 9U.9 0 .0 1.5 0.5 2 3 .2 5.2 h .7 0 .0 U8 .8 0 .0 299.2 0 .0 10.3 6 .0 29.5 7.9 U3.U 0 .0 57.5 76.5 1 1 3 6 .0 16.7 1 5 .6 6.5 37.6 9.1 37.8 0 .0 0 .0 .372.7 0 .0 lli.O 5*6 27.1 6.7 Ul.3 O.U 2 5 .0 U6.7 following the leaf harvest* as is shown by the l a s t column of the table. Both species of c a t t a i l treated with the triethanolamine s a lt of 2 *U-D* on the other hand* did show some regrowth during the eight-week period which followed lea f harvest. This phenomenon is very important in c a t t a i l control for* as already mentioned, between l£ and 20 new c a t t a i l shoots or plant e n titie s have been seen to emerge from basally clipped broad-leaved c a tta il stocks over a period of severai months. II. Root-Immersion Test Results from immersing c a tta il roots for 2b, hours in 1 0 parts per million of systemic herbicidal solution are presented in table 9 . These data show the changes in amount of living material in the above­ ground plant parts brought about five weeks a fte r treatment by the action of certain herbicides placed in contact with the roots through the plant body. Both the ester and s a l t of 2*U-Il were more pronounced in th eir temporary effects on the narrow-leaved c a tta il than was the isopropyl ester of 2*U*$-T. In contrast* the treated broad-leaved c a t t a i l plants showed a much less pronounced difference in temporary effects between 2 , 1i-D and 2 *U*3>-I formulation applications applied in the water immersion solutions. Perhaps the most interestin g resu lts of th is te s t l i e in the fact that* while immersion of narrow— leaved c a tta il roots in 10 parts per million of the various herbicides brought about a pronounced temporary effect* the lastin g r e s u lts , as measured by shoot regrowth eight weeks TABLE 9 EFFECT OF IMMERSING CATTAIL ROOTS FOR 2l* HOURS IN 10 PARTS PER MILLION OF SYSTEMIC HERBICIDAL SOLUTION 5 WEEKS AFTER TREATMENT Plants Treated and Compound Used Narrow-leaved c a tta il Typha angustifolia L. No treatment Shoot Regrowth : Total Leaf Length Total Leaf Weight % Dead 9-22-50 Dead Living OvenOven- Oven- Maximum Leaf Beaker: Dead Living % Fresh dry Fresh dry dry Dead Grams Grams Grams Grams Basis Length cm Number: cm 1-2 : 36.5 332.8 9.9 0.3 0.1 1*.7 1.1 8.3 1*. 2 6-7 :2 1 k .6 115.0 65.1 1.1* 0.8 1.8 0.6 57.1 11.1 2,1*-D, Triethanolamine s a lt 11-12:251.5 82.1 75.1* 1.5 0.7 1.1 0.3 70.0 10.0 2,1*,5-T, Isopropyl ester 16-17:2^.8 376.0 39.1* 1.1* n o 6.9 1.8 33.3 7.2 3-5 :1*62.8 1791*. 8 20.0 6.3 3.8 61.1 12.6 23.2 •8.7 1*53.8 35.6 1*.0 2.1* 19.8 3.7 39.3 0.0 50.9 10.2 36.7 0.0 20.0 0.0 2,1*«D, Isopropyl ester Broad-leaved c a tta il Typha la tif o lia L. No treatment 2,1*-D, Isopropyl ester 8-10:250.7 2,^-D, Triethanolamine s a lt 13-15:589.0 1201,6 32.9 11.3 5.9 2,1*,5-T, Isopropyl ester 18-20:351.2 101*6.1* 25.1 2.1* 1*8.8 3.8 9.6 following leaf harvest, showed no suppression of the new growth. On the other hand, immersion of broad-leaved c a tta il roots in 10 parts per million of similar herbicides brought about a relatively small temporary effect but rather pronounced lasting results as evidenced by no shoot regrowth eight weeks folloying leaf harvest, thus indicating effective suppression of the new growth. I t appears, accordingly, th at the lasting leth al effect on the broad-leaved c a tta il leaves was brought about by transmission of the herbicides through the roots and into the crown of the plants involved. III. Aerial Herbicidal Treatment on Plant Materials Transplanted to Buckets of ^-Gallon Capacity Results from successive clippings of new regrowth developed from the i n i t i a l basally clipped broad-leaved c a tta il root stocks are presented in table 10. These data show certain regrowth changes which take place upon gradual depletion of the carbohydrate root reserves in the i n i t i a l rootstock. Between 15 and 20 new c a tta il e n titie s, capable of independent growth, evolved over a period of several months from one-third of the i n i t i a l basally clipped stocks used in the te s t. In general, with successive clippings, there was (l) a decrease in leaf width of the new plants developed, and ( 2 ) a decrease in the number of visible buds remaining. This study brought out the fact that la te ra l buds are formed 36 w -pOn - o -Cl o O jO j N 5M m • O j O'' n£) OOO' • m Specimen Number ro m m O O^ 00C O HVjJQn co - o M p - NO 0 5 O n O J m m m m \)N O O n O CO Shoot ^ Length M O P- O P- Oo Number of Visible H-* Buds Remaining Average Leaf Width# % Carbohydrate Root Reserve ? I-1 ? O On • • • • • < > 4 38.4 0.9 3 5 56.5 7.0 4 47.6 7 35.6 4 55.9 4 6 8 67.3 9 20.3 12,1 10 55.2 3 11 9.5 4 27.9 27.9 21.6 13 14.0 .. 5.2 8.3 14 7.0 2 5 12 15 5.7 6.4 2.5 77.4 29.1 20.2 11.3 12.2 22.4 67.8 45.3 46.8 38.0 43.1 0.4 36.2 30.9 49.0 51.0 46.1 47.0 34.6 19.2 0.4 44.1 16.5 44.2 59.4 0.4 O O O U"\ 62.9 44.5 7.6 1 1 33.7 7.5 8.5 4.7 52.7 2 0.4 0.3 0.6 3 3 2 26.2 21.0 2.2 8.6 7 .4 0.3 15.5 12.8 3.0 0.2 18.7 0.4 5 40.93 0 35.23 1 46.60 0.5 1 25.0 0.5 6 10.2 0.3 4 49.60 0.3 28.8 25.7 2 0.4 2 0.5 2 0.3 2 37.0 3 0.5 5.7 Dead Dead Dead 6.5 6.0 0.2 2 14.8 14.0 0.4 1 3.8 3.6 0.1 0 32.13 Dead Dead Dead 2 2 Dead Dead Dead 36.5 8 .4 21.0 0.2 1 2 Dead Dead Dead i • 8.0 4 .8 0.2 1 48.77 anew a t a ra te almost commensurate -with new growth. Thus, i t can be expected that a similar phenomenon occurs in the fie ld . Preceding tables show that broad-leaved c a tta il root reserves vary from about 70 percent in the winter dormant condition to a low of approximately 20 percent when they have been depleted by active plant growth. I t can be seen from the la s t column of table U that even after a number of new plants have evolved from the appreciable amount of root reserves remains to carry on oldstock, an newgrowth. Results of the herbicidal action of certain systemic and contact herbicides on narrow- and broad-leaved c a tta il plants three and four weeks after treatment are presented in tables 11 and 12 respectively. The contact herbicides, modified rosin primary amine, pentachlorophenol and sodium trichloroacetate, applied 2 b hours after approxi­ mately pounds of systemic herbicide had been applied to the same plants, appeared to p a rtia lly nullify some of the effects usually associated with systemic herbicides. On the narrow— leaved c a tta il 2^.30 pounds, and on the broadleaved c a tta il U1 .U6 pounds, 2 ,U—0 acid equivalent per acre, were effective in lowering the carbohydrate root reserve to 2JU.77 and 2 U.2 0 percent, respectively, and in both Instances this relative carbohydrate root exhaustion (see tables 3 — 7 ) effected complete eradication. lb- treated plants in similar stage of growth had a carbohydrate root reserve of approximately 60 percent for both species of c a tta il. I t was interesting to note that the plants treated July 7> and reported in table 1 1 , showed less effects from the systemic herbicide 37 TABLE 11 HERBICIDAL EFFECT OF CERTAIN SYSTEMIC AND CONTACT HERBICIDES ON NARROW - AND BROAD-LEAVED CATTAIL PLANTS THREE W EEKS AFTER TREATMENT Total Leaf Length : Plants Treated and Compound Used Narrow-leaved c a tta il Typha angustifolia L. No treatment 2 ,U-D, Isopropyl ester 2 ,U-D, Triethanolamine s a lt 2 , U , 5 -T * Isopropyl ester 2 ,U-D, Na s a lt 2 ,U-D, Na s a lt 2 ,U-D, Na s a lt 2 ,U-D, Na s a lt -^-Modified rosin, primary amine 2,U-D, Na s a lt -*Pentachlorophenol 2,1|-D, Na s a lt -“'Trichloroacetate, Na s a lt Broad— leaved c a tta il Typha la tif o lia L. No t r eatment 2 ,U— D, Isopropyl ester 2 ,U-D, Triethanolamine sa lt 2 ,U ,£ - T , Isopropyl ester 2 ,U-Q> Na s a lt 2 ,U-D, Na s a lt 2 ,U— D, Na s a lt 2 ,U— D, Na s a lt -^Modified rosin, primary amine 2 ,U-D, Na s a lt -“-Pentachlorophenol 2 ,U-*D, Na s a lt -*Trichloroacetate, Na sa lt Found.s per Buck et Acre Number 1 8 .5 8 2 8 .3 5 9.U7 6 .6 7 1 1 . 2k 2 5 .3 0 lit. .0 6 3 3 5 .0 1 1 .9 5 1 9 .SU 2 1 .0 9 3 8 8 .2 2 U.0 U U5 -3 7 3 3 .5 1 1 1 .2 3 1 9 .6 8 U1 .U6 1 7 .5 7 U1 8 .7 1 8 .9 0 3 1 .5 1 1 8 .9 8 3 U9 .U Dead cm Living cm /3 Dead 1056 1051 1052 1053 105 u 1055 1057 0 .0 7 0 1 .7 1 U 1 .1 6U .2 8 0 .0 7 2 1 .u 7 5 1 .2 1058 3U 2 .0 1 6 6 .0 6 7 .3 1059 3 0 5 .7 U 16.9 U 2.3 1060 8 2 9 .2 3 5 1 .3 7 0 .2 1006 1001 1002 1003 100U 1007 U 38.5 U 3 5 .0 7 7 3 .6 2 2 7 .1 5 7 6 .6 1075.6 8 0 0 .9 2 6 5 9 .6 1 U.2 1 1 5 6 .0 2 3 .0 7 9 8 .2 U9 . 2 2 5 1 2 .5 8 .3 7 1 3 .5 UU.7 7 2 0 .2 5 9 .9 0 . 0 1 0 0 .0 1008 UU8 . 3 1 5 2 9 .3 U2.9 1009 5 3 5 .0 1 6 3 8 .1 2 U.6 1010 1332.2 105U .9 5 5 .8 ioc5 1 0 8 7 .6 0 .0 10U 1.U U0 .3 1 U5 2 .3 8 .9 2 0 3 1 .2 3 .1 U2 8 .6 1 5 .7 7 8 .2 2 0 0 .7 0 .0 1 0 0 .0 -“-Contact herbicide applied 2 k hours a lte r preceding systemic herbicide v/as applied. Total Leaf Weight Dead Living Oven: OvenFresh dry Fresh : dry Grams Grams Grams : Grams 0.0 8.9 1.9 1.0 0.9 12.3 9.2 G.O 5.1 1.2 0.2 o.5 U.8 5.7 h .l 28 .8 s % Dead Ovendry Basis : : : : : : 7.1 6.0 13.6 12.9 5.0 0.6 0.0 0.0 5o.U 8.1 1.6 9.1 38.9 100.0 3.0 2.0 : 0.3 7.U 1.8 12.5 : 9.9 6.5 11.3 : 13.1 7.8 16.3 5.9 10.9 20.9 13 . h 9.3 5.3 13.1 3.2 7.0 15.9 11.5 89.3 21;.6 21.1 98.7 2li.5 25.3 0.0 2lu5 Initial Individual Shoot Height above H gO Surface cm Number of Green Shoots Developing after Treat­ ment Percentage Carbohydrate Root Reserve after Treat­ ment Measured Percentage Survival 67^9 71;,60,58 95,77,79 6 8 , 8)4, 6 5 , 61' 86 7U, 78,1;8,52 7U,5i,70,75 1 5 3 6 0 2 0 0 / V^ /o 81,7U 1 21 2.1; U2.8 60,69 0 58 2.U 73.0 82 , 81;, 79 2 28 : : : : : : : 25.9 6.8 5.3 29.8 7.5 7.6 0.0 26. L ; U3.8 81,91,72,86 59,75 7h,99 91,105 78,72 86,89,95 6 3 2 h 100.0 86,78 0 16.7 52.3 : 16.5 5 0 .3 8)4 , 88,103 5 53 12.8 7.9 53.5 : i5.lt 33.9 9 1 , 8)4 rn 71 21.9 19.3 28.7 : 6.2 75.7 91,72,102 a 3U 23.5 51.U 51.8 19.0 3.6 0.0 71.2 9.7 U8.3 6 7 .6 100 55 92 2h,77 98 88 16 0 80 67 ho 91 53 36 - 2 2 2U.20 0 applied than did similar jplants treated August 25, and reported in table 1 2 • This can be accounted for p a rtia lly , perhaps, by the fact that cooler temperatures prevailed immediately following the herbicidal application on the second group of c a tta il plants and thus there was less stimulation for new growth in the la tte r group. The arithemetical average of the daily maximum, minimum and 8 a.m. temperatures for the two periods was as follows: August 2b to September 22 July 7 to July 28 maximum 82.3 minimum 51-5 8 a.m. 68. k maximum 8 0 .0 minimum h k .6 8 a.m. 6 2 .0 Measurement of percentage survival at the termination of the te s t is presented in tables 11 and 1 2 . Because of temperature differences during the two series of tests the comparative value of measured percentage survival of the ca tta ils three weeks aftex' the f i r s t te s t and four weeks after the second te st remains uncertain. In the instance Yfhere 3*92 pounds of 2,U-D acid equivalent ap­ peared to bring about a 100 percent k i l l in the narrow-leaved cattails chemical analysis showed a carbohydrate root reserve of approximately U2 percent. This was one of the few instances in which Vatsol 0T-B was used as a wetting agent. XV. Aerial Herbicidal Treatment on Plant Materials Transplanted to Metallic Containers of 27-1/2 Gallon Capacity 38 TABLE 12 .HERBICIDAL EFFECT OF CERTAIN SISTEMIC AND CONTACT HERBICIDES ON NARROW- AND BROAD-LEAVED CATTAIL PLANTS FOUR WEEKS AFTER TREATMENT Total Leaf Length Plants Treated and Compound Used Narrow-leaved c a tta il Typha an eustifolia L. No t r eatment 2 ,i*-D, Isopropyl ester 2>l*-D> Triethanolamine s a l t Pounds per Bucket Acre Number Dead cm Living cm % Dead 26.95 3 2 .2 8 1066 7 2 8 .0 1061 860.3 1 0 6 2 2351*.7 1159.8 699.5 1729.6 3 8 .6 55.2 57.7 2,1*, 5-T* Isopropyl ester 2,1*— D, Triethanolamine s a l t 19.31 12.65 1063 595.7 1061 * 9 6 3 .0 933.3 121*3.3 39-0 1*3.6 2,1*-D, Na s a lt 2,1*-D, Na s a lt 3.92 13.69 1065 781 *. 1 1067 2 0 5 1 .0 0 .0 1289.7 1 0 0 .0 6 1 . 1* 23-55 1*0 . 0 6 7 1 2 .0 1 0 6 8 318.3 1069 651*. 7 1 0 7 0 2 6 3 U. 2 51*6.3 1*07.5 151*.7 3 6 .8 6 1 .6 91*.5 1016 1011 1012 675.5 958.3 920.1 916.5 721*.7 8 0 0 .3 1*2 . 1* 56.9 53.3 1013 926.2 653.7 5 8 .6 2,1*~D, Na s a lt 2 I4.— D, Na s a lt Pentachlorophenol Broad-leaved c a tta il Typha l a t i f o l i a L. N0 t r eatment 2,1*-D, Isopropyl ester 2,1*— D, Triethanolamine s a lt 14*. 80 37.08 2,1*,5-T, Isopropyl ester 2,1*-D, Triethanolamine s a lt ^ 2 ,U-*D, Isopropyl ester 2,1*~D, Na s a lt 2,1*— D, Na s a lt 2,1*-D, Na s a lt 2,1*-D, Na s a lt Pentachlorophenol 29.87 15.71 13.11 11*. 32 17.55 20.75 61.15 571.9 1011 * 9 8 1 .8 1015 1769 . 2 1017 975.1 1 0 1 8 1*20.9 1019 530.8 1 0 2 0 520.9 1*1*1 . 1* 6 9 . 0 1 0 0 .0 0 .0 1*1*. 6 1 2 1 3 .3 7 2 2 . 1* 3 6 .8 39.2 823.9 83.9 99.7 *-A mixture of the ester and preceding s a l t of 2,1*-D were used. Total Leaf Weight Dead Living OvenOvenFresh dry Fresh dry Gram s Grams Grams Grains I4.O 27.0 1*.8 II1.6 1 8 .1 10.2 8 .0 6U-3 21.3 3.3 7.2 31.6 Ho.5 10.2 12 oil 13.7 7c3 U2 «U 6.5 19.5 0 .0 67 .8 22.9 2.7 6.7 1*2.3 2.1 1*.7 11.7 11.5 1*.5 Ii.5 23.0 1 .0 0 .6 21.6 7.7 9.3 22.7 10.8 ii3.9 25.7 33.0 1 5 .8 1 0 .1 1 1 .5 12.1 7.0 25.0 7.2 20.2 13.7 37.1 19.5 7.8 0 .0 70 .U 35 .6 31.5 0 .0 5.1 5.3 23.8 l*-5 15 .1* 85.7 53.9 13.9 12.5 16.0 22.1 7.U Ii.7 9.0 1.3 0 .0 2l*.5 12.lt 9. k 0.2 % Dead Ovendry Basis 28.2 37.5 1*0.7 2li.ii 3 I4.I 1 100.0 1*6 .0 3 1 .8 51.1 97.5 3 2 .8 Initial Individual Shoot Height above H ^O Surface cm 90,83,62,115,110 81 , 118,116 120 , 71 , 131 , 11*1 , 108 , 130,90,153 121*, 100,130 51,93,11*2,99,116, 119 112,83 112 , 11*2 , 108 , 168 , 11*3 89 , 1*5 ,8 0 102,108,87 135,125,129,113, 91,81* 1 0 1 121 , 22,110 1*7.9 109,86,83,27 1*8 . 1* 77,112,85,56,116, 87 ' 1*9.3 83,91*,95,111*,76 116,92,120,99 100.0 136,153,112,133 1*7.1* 150,153,139 37.1* 120,90,86 33.3 90,23,90,70 97.8 126,79 63.7 Number of Green Shoots Developing after Treat­ ment Percentage Carbohydrate Root Reserve after Treat­ ment 58.67 Measured Percentage Survival 67 51* 5i 0 1 60.80 68 61 0 1 1*2.13 65.63 0 1*6 I* 60.30 66 1*1* 1* 0 0 0 56.77 0 0 1*9 1*6 0 1 0 0 0 0 3 62 1*8 56.53 52.50 59.20 5U.97 31* 0 51* 63 /1 61* 9 The submerged aquatic plants treated, systemic and contact herb­ icides used, th eir ra te s, in pounds per acre acid equivalent basis, and estimated percentage survival are presented in table 1 3 . At the conclusion of the experiment the so il in the bottom of the tanks was examined for the presence of living underground propagules and from this physical examination the following observations were made on the treated waterweeds: True Waterweed With one treatment. 5>»09 pounds of 2 ,U~L effected complete eradication. 8.U2 pounds 2,1|,5>--T effected complete eradication. With tvj o treatments. 11 pounds 2,U,3“T, followed seven weeks la te r by 9 pounds 2 , 1;, 5 — T, effected complete eradication. Leafy Pondweed With one treatment. 5 pounds of 2 ,U-L effected complete eradication. With two treatments. 5>.5> pounds of 2 ,U-D followed five weeks la te r by 3 .£ pounds 2 ,U~L effected complete eradication. 79 pounds of anhydrous copper sulphate followed three weeks later by 116 pounds of anhydrous copper sulphate showed no visible control or eradication effects seven weeks after second treatment. American Pondweed With one treatment. 6 pounds of 2,U-D effected good control with medium infestations. 39 TABLE 13 AM OUNT, IN POUNDS PER ACRE, OF PHENOXYACETIC ACID AND OTHER COM POUNDS USED O N THE AQUATIC AND M ARSH PLANTS, LISTED TOGETHER WITH THE ESTIMATION OF PERCENTAGE OF SURVIVAL AT THE TERMINATION OF TEST Pounds per Compound : Acre Acid Equivalent 2,1-D, Sodium sa lt True waterweed : 32.36 2,1-D, Sodium sa lt Anacharis canadensis i 21.73 2,1,5-T, Isopropyl ester (Michx. P1anchon 1 9 .9 k 2,1-D, Triethanolamine sa lt 13.62 2,1-D, Triethanolamine sa lt .. 13-10 2.1-D, Isopropyl ester 12 .I|2 2.1-D, Triethanolamine sa lt 11.61 2,1-D, Isopropyl ester 2,1-D, Triethanolamine s a lt 11.30 6.1-2 2,1,5-T, Isopropyl ester 2,1-D, Sodium s a lt 5.09 Leafy pondweed 195.01 Anhydrous copper sulfate 2,1-D, Sodium salt Potamogeton foliosus 21.87 Raf. 2;1-D, Triethanolamine sa lt l l .8 0 2,1-D, Triethanolamine salt 13.51 2,1,5-T, isopropyl ester 11.90 2,1— D, Isopropyl ester 1 1 .1 8 2,1-D, Isopropyl ester . . 10 .35 9.21 .. 2,1-D, Isopropyl ester 2,1.5-T, Isopropyl ester 8.18 2 ,1-D, Sodium sa lt 6 .8 7 2 , 1-D, Triethanolamine sa lt 6.72 2,1-D, Isopropyl ester 5 .6 5 2,1— D, Sodium s a lt 1.93 2,1,5-T, Isopropyl ester 3.19 Alpha benzene hexachloride American pondweed 1277.62 Anhydrous copper sulfate Potamogeton nodosus 191.99 2,1,5-T, Isopropyl ester . 1 6 .8 0 Poiret 2,1-D, Triethanolamine s a lt 13.62 2 ,1-D, Isopropyl ester 11.76 2 ,1-D, Triethanolamine salt 1 1 . 1+0 2 ,1-D, Isopropyl ester 9.36 2,1,5-T,- Isopropyl ester 9.21 2,1-D, Sodium salt 37.60 Gigantic sago pondweed 2 ,1-D, Triethanolamine salt 22.18 Potamogeton pectinatus 2,1-D, Sodium salt L. 21.58 2,1-D, Triethanolamine sa lt 20.37 Plants Treated Estimated Percent Survival 0 0 0 100 0 30 15 0 0 0 100 0 0 0 0 0 0 0 0 0 0 5. 0 85 . _ . 5 10 . ...... 5 2 .........1 5 5 60 5 %_______ , . 5 ------. . j L . - TABLE 1 3 —• C o n tin u e d Pounds per • Estimated Plants Treated Acre Acic : Compound Percent Equivalent * Survival Gigantic sago pondweed .... 17.52 :2.L.5-T. Isopropvl ester 5 Potamogeton pectinatus 16 .6 1 : 2 ,l4— D, Isopropyl ester 10 L.—Continued 15 .66 :2,U— D* Isopropyl ester 10 . . . 13.62 :2,ij.-D9 Triethanolamine s a lt 5 13.53 . :2»1;-D, Triethanolamine s a lt . . . 3.0. . : 2 »1j.Xd, Isopropyl ester 20 11.37 :2,ij.-D. Triethanolamine s a lt 2 - ...11.35 20 ..... 11.30 :2,U-D» Isopropyl ester 10. 8§ :29L-D9 Isopropyl ester 1 10.62 20 : 2 ,h ,5 ~ T s Isopropyl ester...... 1 0 .0 8 r2,iu5-T. Isopropyl ester 75_ . 8.83 t2,k,5-T, Isopropyl ester ....... 85 8.11 s2,Lu5— 2 1 T, Isopropyl ester 5 .65 t 2 ,U-D, Is opropyl ester . 70 :2 ,L-D, Sodium s a lt . i5_ . Slender sago pondweed : Sodium arsenite 5>17.lh 25 . Potamogeton pectinatus _3 3 1 . & __ :Anhydrous copper sulfate 100 L. 1 00 ; • Anhydrous copper sulfate 1 9 6 .5 9 . Richardson* s pondweed D, Triethanolamine s a lt 27.12 :2,h— i 5 .. . . 10 Potamogeton richardsarw 22.57 :2,U.-D, Isopropyl ester : 2 ,U-D, Isopropyl ester i i (Ar. Benn#) Rydb* . 1 5 .0 7 85. ... 2 11.26 : 2 ,U— D, Triethanolamine s a lt 100 :2,I|.-.5-T, Isopropyl ester 5 .8 8 0 Horned pondweed 806.3& :Alpha benzene hexachloride 1 00 :Anhydrous copper sulfate 1X6 .9 8 Zannichellia p alu stris uo 356 .1 2 : Sodium arsenite L0 0 328.30 :TCA, Sodium s a lt 0 : 2 ,L iX d, Sodium s a lt 21.87 0 D. Isopropyl ester 21.53 :2.U— 20 D, Sodium s a lt 21.51 0 13.53 :2,li-.b, Triethanolamine s a lt 30 D, Triethanolamine s a lt 13.30 :2 9ij— -:a B, Triethanolamine s a lt 12 .U2 : 2 ,U— 0 Isopropyl ester 10 11.3L :2.L-D. Isopropyl ester 0 10.77 r 2 .ij.-D, Isopropyl ester 50 . 8.72 : 2 ,lw^-T, Isopropyl ester 0 2 .U.5-T. Isopropyl ester, 7 .2 0 parts of isopropyl ester and trie tk X ■if-Pounds consist 5 O ■snolamine s a lt. 9.5 pounds of 2,lj.-D e f f e c t e d good control with heavy in festa tio n s. With two treatm ents. 5 . 5 pounds of 2 , I4—D followed five weeks l a t e r by L j. pounds of 2,1|-D effected 95 percent eradication. 1 3 3 pounds of anhydrous copper sulphate followed five weeks la te r by 35 9 pounds of anhydrous copper sulphate effected 6 0 percent control or eradication seven weeks a fte r second treatment* 7 9 2 pounds of alpha benzene hexachloride followed five weeks la te r by I486 pounds of alpha benzene hexachloride effected 95 percent control or eradication seven weeks a fte r second treatment. Gigantic Sago Pondweed With one treatment* 1 1 pounds of eith er 2 aU— D or 2 , 14, 5 - 1 effected good c o n h o l. With two treatm ents. 6 pounds of 2 yh—B followed seven weeks l a t e r by 5 pounds of 2 ,U— D effected 9 9 percent eradication. 8 pounds of 2,U,5-T followed one day la t e r by U3U pounds of modified rosin primary amine effected 9 8 percent eradication. With three treatm ents. 1 3 . 5 pounds of 2 ,Li~D followed one day l a te r by 820 pounds of alpha benzene hexachloride, followed six weeks la te r by 5 U7 pounds of alpha benzene hexachloride effected 9 5 percent eradication. 1 1 . 5 pounds of 2 ,U— D followed one day l a te r by 820 pounds of sodium chlorate, followed six weeks la te r by 8 2 0 pounds of sodium chlorate effected 8 0 percent eradication. 11 pounds of 2 ,U-i) followed one day la te r by 3 7 0 pounds of trich lo ro acetate, followed six weeks l a te r by 10.5 pounds of 2 ,iL|.— D effected 95 percent eradication. 16 pounds of 2 ,U-*h followed one day la te r by 6 I48 gallons of lylene and 6.5 gallons of California Spray Ortho no. 5^ followed six weeks l a t e r by 21.5 pounds of 2 , 14— D effected 95 percent eradication. Uo S l e n d e r S ago P o n d w ee d W ith tw o t r e a t m e n t s . 152 pounds of anhydrous copper sulphate followed three weeks la te r by 182 pounds of anhydrous copper sulphate effected no v isib le control or eradication seven weeks a fte r second treatment. 2 U2 pounds sodium arsenite followed four weeks la te r by 275 pounds sodium arsenite effected no v isib le control or eradication seven weeks a fter second treatment. Richardson1s Pondweed With one treatment. 22,5 pounds of 2 ,1 ;D effected 90 percent eradication. With two treatments. 7.0 pounds of 2,1;— D followed seven weeks la te r by 1;.J? pounds of 2,1;— D effected 9 8 percent eradication. The triethanolamine s a lt of 2,U-i) was appreciably more effective than i t s isopropyl ester in bringing about Richardson’s pondweed k ill. Horned Pondweed With one treatment. 7.2 pounds of 2,1;,5~T effected complete eradication. 806 pounds of alpha benzene hexachloride, 638 gallons of xylene, 6.5 gallons of California Spray Ortho no. 5 effected complete eradication. With two treatments. 6 .0 pounds of 2 , 1;— D followed seven weeks la te r by 5 « 0 pounds of 2 , 1;— D effected complete eradication. 328 pounds of trichloroacetate followed 12 weeks la te r by 356 pounds of pentachlorophenol effected complete eradication. l5 l pounds of sodium arsenite followed four weeks la te r by 205 pounds of sodium arsenite effected 60 percent eradication. U1 216 pounds of anhydrous copper sulfate folio-wed four -weeks la te r by 231 pounds of anhydrous copper sulphate effected no visible control or eradication seven -weeks after second treatment. Phenoxyacetic acid treatments followed 2 U hours la te r with contact herbicides seemed to be less effective in control or eradication than the phenoxyacetic acid treatment without any following contact herbicidal treatment. Systemic herbicidal applications made in repeat treatments appeared to be more effective than when the same quantity was applied as a single treatment. In general, the following physical phenomena were observed for the phenoxyacetic acid treated submerged aquatic vascular plants: One week after treatment stems and leaves showed pro­ nounced curvatures. Four weeks after treatment chlorophyll depletion was apparent with most leaves and stems brown, and apparently dead. Six weeks after treatment regrowth, i f any, developed from laten t propagules, e .g ., buds, seeds, etc. I t was interesting to note th at some survival occurred in a ll of the treated tanks containing American, gigantic sa.go, and Richardson* s pondweeds, even though large quantities of systemic herbicides were us ed. I t has been observed by Moore ( 2 2 ) that when tubers o ccur in twos, as is the case with Potamogeton pectinatus, only the larger one develops the shoot. becomes detached. The smaller tuber does not sprout unless i t In th at case only, i t develops an individual plant. Potamogeton nodosus accomplishes vegetative propagation also by subterranean scaly buds which generally grow in pairs a t the end of the rootstock. Potamogeton pectinatus seeds were gathered July 12, 1 9 k 9 , from the Federal Center lake a t Denver, Colorado, and kept in cold storage through the winter. On April 19, 1950, 100 seeds were placed in a covered 2 -ounce ja r containing tap water, which in turn was placed in the greenhouse to observe germination. Germination proceeded as follows: Number of Germinated Seeds Removed from Jar Germinati on Date 18 U -2 5 -5 0 5-1-50 5- 5 -5 0 2 3 5-9-5o 2 2 2 1 0 0 3 5 - i 2- 5o 5- 1 5 -5 0 5 - 2 2 -5 0 6 - 2 -5 0 6 - 1 6 -5 0 7-1 0 -5 0 Apparently, the systemic herbicides did k i l l the above-ground vegetative growth present a t the time of tank treatments. The per­ centage survival, present a t the termination of the experiment, appears to have come from la te n t bud propagules or seeds which had not started growth a t the time of a erial herbicidal treatment. This work suggests a new, effective and economically practical technique for applying phenoxyacetic acid compounds on submerged waterweeds growing in irrig a tio n ditches• the following steps: This method consists of (l) shut off irrig a tio n water flow and allow maximum possible gravity drainage to obtain in ditch bottom; ( 2 ) apply aerial herbicidal spray of 2 ,U-D or 2,U,5-T, using a wetting U3 agent, on the unsupported waterweeds lying in the bottom of the gravity drained ditch; ( 3 ) allow two to six hours for absorption of chemical by water plants; and (U) turn water back into irrigation ditch. V. Miscellaneous Tests Detection of 2,l|.-D in roots of top-treated c a tta il plants. In the te s t performed to determine presence of 2,1<-D in the roots of top-treated c a tta il plants, 9 out of II4. of the c a tta il roots examined gave the color reaction which indicated presence of 2,U-D. This indicates that the waterline as such does not act as a barrier to the passage of 2,lj-D through c a tta il tissues. Effect of systemic herbicidal applications on water sedge. Results showing depletion of carbohydrate root reserves as well as leaf killing effect in water sedge plants treated with different systemic herbicides are presented in table II4.. TABLE 1U DEPLETION OF CARBOHYDRATE ROOT RESERVES IN W ATER SEDGE PLANTS TREATED WITH DIFFERENT SYSTEMIC HERBICIDES Herbicidal Compound Pounds per Acre Acid Equivalent Pex-centage Carbohy­ drate 7 Weeks Fol­ lowing Treatment No treatment 2,U-D, Isopropyl ester and -^Triethanolamine sa lt U5 .8 7 3 5 .5 2 2U.30 Measured Percentage Survival 100 8 2,U,9-T, Isopropyl ■ 21 ester 1 ^ .6 8 2U.U7 . . n S t -x-Applied approximately in the proportion of o parts sa lt to 1 part ester. hh Systemic herbicidal application in both instances decreased the carbohydrate root reserve by approximately j?0 percent and effected an approximate 90 and 80 percent k i l l "when using 2 ,U-b a t a rate of 3!?»£2 pounds and 2 ,Hj!?-,T a t a rate of l4.L 1.-68 pounds acid equivalent per acre, respectively. This -work indicates th at systemic herbicidal treatments on -water sedge plants can be used to effectively deplete the carbohydrate root reserves and thereby bring about a permanent k i l l of the treated plants. SUMMARY In order to grow diversified crops on much of the land in the western United States i t is necessary to supplant the moisture provided by nature in the form of ra in , snow, e tc ., with additional water which reaches farm lands through established irrig atio n canal distribution systems. These systems frequently support heavy growth of vascular plants which prevent or slow down the passage of water. Since reducing the carrying capacity of a canal makes i t necessary to deprive some potentially crop-producing land of the required water to bring the crop to a satisfactory harvest, i t is imperative to keep the waterways open. This thesis is a report of certain pertinent investigations which suggest improved and more effective fie ld techniques to accom­ plish a solution of the problem a t hand. In the le a f-tip immersion te s t the isopropyl ester of 2,U,5-T and 2 ,l|.-D were more pronounced in their lethal effects, both temporary and lasting, on the narrow-leaved c a tta il than was the triethanolamine s a lt of 2 , 1+-D. Lethal effects of the ester of 2 ,U-D appeared to be transmitted through the immersed leaf tip past the waterline, and into the crown of the c a tta il plants. Associated with this phenomenon is the fact that no shoot regrowth developed in the ester treated plants. The immersion of broad-leaved c a tta il roots in 10 parts per U6 million of the s a lt and ester formulations of 2,li-D and the ester formulation of 2,1;,5>~T, brought about lasting resu lts evidenced byno shoot regrowth eight -weeks follo-wing le a f harvest. I t is a common occurrence for basally clipped broad-leaved c a tta il rootstocks placed in beakers of water each to evolve between If? and 20 new c a tta il e n titie s capable of independent growth. Contact herbicides on c a tta ils , applied 21; hours after approxi­ mately 15 pounds of systemic herbicide had been applied, appeared to n ullify the more lasting effects of the systemic herbicides. Systemic herbicidal applications made in repeat treatments appeared to be more effective than when the same quantity was applied as a single tr e a t­ ment. On the narrow-leaved c a tta il 2]?.30 pounds, and on the broad­ leaved c a tta il I4I.I 4.6 pounds, acid equivalent, per acre were effective in lowering the carbohydrate root reserve to 2 1;.77 arid 21;.20 per­ cent, respectively. Untreated plants in a similar stage of growth had a carbohydrate root reserve of approximately 60 percent for both species of c a t t a il . When aerial herbicidal treatments were made the following single or repeated 2 ,U**U applications were effective in obtaining complete, or nearly complete, eradication of the waterweeds growing in the so il bottom of the treated tanks. U7 Name of Plant Narrow-leaved C attail Broad— leaved C attail True Waterweed Water Sedge Leafy Pondweed American Pondweed Gigantic Sago Pondweed Richardsont s Pondweed Horned Pondweed Pounds per Acre Number of Tr eatments Percent Eradication 29 Ul 1 1 1 1 1 1 2 100 27 5 10 20 12 11 2 2 100 100 89 100 99 9$ 98 100 I t appears th a t phenoxyacetic acid derivative applications on submerged aquatic plants should be made before the plants reach the fru itin g stage. Broad- and narrow-leaved c a t t a i l roots showed considerable variation in the amount of carbohydrates present during the growing season. Highest carbohydrate was present during the winter dormancy period and lowest carbohydra.te was associated with production and maturation of male and female fru itin g bodies. Without exception i t can be said th at root reserves are low from the time of the f i r s t appearance of the fru itin g stocks u n til pollination has been completed. However, in some c a tta il growing s ite s fru itin g bodies are entirely lacking and, accordingly, other above-ground phenomenon, associated with low root reserves, must be used, to determine optimum time for treatment to obtain maximum c a tta il control or possible eradication with minimum time, effo rt, and cost. C attail growth in the water inundated, growing site s began when the rising spring water temperature reached approximately U6 C F. In the narrow-leaved and broad-leaved c a tta il growth site s not U8 inundated by ■water root reserves were a t a minimum when the plants had attained a height cf approximately 1 0 0 -1 3 0 cm and ^0 - 1 2 0 cm above the ground lin e respectively. In general, for both c a tta il species the period of seed stalk elongation and rapid expansion of the female spike was one of low carbohydrate root reserves. Both species of c a tta ils attained maximum leaf width immediately preceding th e ir respective periods of low carbohydrate root reserve. In a l l five c a tta il growth situations investigated there was a marked tendency for percentage dry matter in roots to be relativ ely high when percentage carbohydrate in roots was high and similarly to be low when percentage carbohydrate was low. In contrast to the roots, the percentage dry matter in the shoots, in the five c a tta il situations investigated, increased apparently a.t the expense of the carbohydrate root reserves which at the same time were being depleted. The chromotropic acid te s t indicated 2,b.-D to be present i':i the roots of 9 of 1 U c a tta il samples taken from plants receiving aerial applications of 2 ,Li-D. This indicates that the waterline is not a b arrier to the passage of 2 ,U-D through the vascular tissue of aquatic p la n ts. w LITERATURE CITED 1. 2 . Balcom, R. B. Control of Weeds on Irrigation Systems. United States Department of In terio r, Bureau of Reclamation, Washington, lUo pp. 1914-9. Bohmont, D. W. Using 2 ,l_i-D in Wyoming. Wyoming Agricultural Experiment Station, Laramie, Bull. 291,- 19li9. 3. Corrington, J. D. Working with the Microscope. Company, New York, 1+18 pp. 19Ul. McGraw-Hill Book Uc Coulter, J. M., and A. Nelson. New Manual of Botany of the Central Rocky Mountains. American Book Company, New York, 61+6 pp. 1909. 3. Crafts, A. S. Control of aquatic and ditchbank weeds. California Agricultural Extension Service, Davis, Circ. 158, 19h9° 6. Eames, A., and L. H. MacDaniels. An Introduction to Plant Anatomy. McGraw-Hill Book Company, New York, 309-31U. 1923'. . Mitchell, and R. W. Heinen. Using 2,1).— D safelyQ 7o Evans, L. S ., J. W United States Department of Agriculture, Washington, Farm. Bull. 200 £, I 9 I48. 8. Fassett, N. C. A Manual of Aquatic Plants. Company, New York, 382 pp. 19U0. 9. Fernald, M. L. The linear-leaved North American species of Potamogeton, Section Axillares, Mem. Amer. Acad. Arts & S ci., 17(1)-.1-183. 1932. . Fernald, M. L. Gray’s Manual of Botany, ed. 8, American Book Company, New York, 1632 pp. 195>0. . Freed, V. H. Qualitative reaction for 2,U-dichlorophenoxyacetic acid. Science. 107: 98-99, 19U8. . Freeland, R. 0. Effects of 2,U-D and other growth substances on photosynthesis and respiration in Anacharis. Bot. Gas.Ill:319— 32U, 195>0. 10 11 12 McGraw-Hill Book 13. Grigsby, B. H., B. R. Churchill, C. L. Hamner, and R. F. Carlson, Chemical weed control. Michigan Agricultural Experiment Station, East Lansing, Circ* Bull. 21 I4., 19U9» ll+o Hall, T. F ., and A. D„ Hess. Studies on the use of 2,1+-D for the control of plants in a malaria control program. J. Nat. Mai. Soc. 6 : 99-116, 191+7. 15 o Hotchkiss, N., and H. L. Dozier. Taxonomy and d istrib u tio n of N. American c a t - t a i l s . Amer. Midland N atu ralist. 1+1(1): 237-251;, 191+9. 16 . Lange, N. A. Handbook of Chemistry. Sandusky, Ohio, 1232-1239• 19Uho 17. Lepper, H. A., (Chairman). O fficial and Tentative Methods of Anal­ ysis of the Association of O fficial Agricultural Chemists. As­ sociation of O fficial Agricultural Chemists, Washington, 932 pp. 191+5. 18. Loomis, W. E., and C. A. Shull. Methods in Plant Physiology. McGraw-Hill Book Company, New York, 1+72 pp. 1937. 19. M iller, E. C. Plant Physiology. York, 776. 1938. Handbook Publishers, Inc., MeGraw— Hi11 Book Company, New 20 . M itchell, J . W., and J. W. Brown. Effect of 2,l+-dichlorophenoxyacetic acid on the read ily available carbohydrate constituents in annual Morning-glory. Bot. Gaz. 107: 120-129, 191+5. 21 . __________________ , and _________________ . Movement of 2, ]+-dichlorophenoxyacetic acid stimulus and i t s re la tio n to the translocation of organic food materials in p lan ts. Bot. Gaz. 107: 393— 1+07* 191+6. 22 . Moore, E. The potamogetons in re la tio n to pond cu ltu re. United States Department of Commerce, Bureau of F isheries, Washington, Bull. 33, 1915. 23. Muenscher, W. C. Aquatic Plants of the United S tates. Comstock Publishing Company, In c ., Ithaca, New York, 371+ PP° 191+1+. 2 l+. Ogden, E. C. The broad— leaved species of Potamogeton of North America north of Mexico. Rhodora. 1+5: 57— 105, 109— 163, 171— 2ll+, 191+3. 25. Rasmussen, L. W. The physiological action of 2 ,l+-dichlorophenoxyacetic acid on dandelion, Taraxacum o ffic in a le . Plant Physiol. 22: 377-392, 191+7. 26. Robinson, B. L., and M. L. Fernald. Gray‘s New Manual of Botany, ed, 7, American Book Company, New York, 926 pp. 1908. 27. Rydberg, P. A. Flora of the Rocky Mountains and Adjacent Plains„ P. A. Rydberg, New York, 1110 ppD1917. 280 St. John, H. A revision of the North American species of Potamogeton of the Section Coleophylli. Rhodora. 18: 121-138, 1916. 29. Sass, Jo E0 Elements of Botanical Microtechnique. Book Company, New York, 222 pp. 191+0. 30. Schmeil, 0o, and J. Fitschenc Flora von Deutschlando Quelle and Meyer Publishing Company, Leipzig, Germans’-, J439 pp„ 1916. McGraw-Hill 31. Schaffer, P. A», and A. Fc Hartmann. Ihe iodometric determina­ tion of copper and i t s \ise in sugar analysis. J. Biol. Chem. 1+3: 3 6 5 - 3 7 3 , 1 9 2 0 - 2 1 . 32. Smith, F. G., Co L. Hamner, and R. F. Carlson. Changes in food reserves and respiratory capacity of bindweed tissues accompany­ ing herbicidal action of 2,I)-dichlorophenoxyacetic acid. Plant Physiol. 22: 58-65, 19U7. 33. Speirs, J. M. Summary of lite ra tu re on aquatic weed control. Fish Cult. 3(1+) s 20— 32, 191+8. 3l+o Surber, E. W«, C. E. Minarik, and W. B. Ennis, Jr. The control of aquatic plants with phenoxyacetic compounds. Progressive Fish-Culturist. 9(7):ll+8, 191+7. 35* Thimann, K. V. Use of 2,l+-dichlorophenoxyacetic acid herbicides on some woody tropical plants. Bot. Gaz. 109: 33l+— 3l+0, 191+8< , 36. Weaver, R. J . , and H. R. DeRose. Absorption and translocation of 2,1+— dichlorophenoxyacetic acid, Bot. Gaz. 107: 509— 521, 191+6. 37. Woodman, A. G. 607 pp. 191+1. Food Analysis. Can. McGraw-Hill Book Company, New York, XTCIMHddV Side view Aerial view Plate I . Arrangement for immersion of broad- and narrow-1eaved c a tta il leaf tips (one tip per plant) in jars of commercial herbicidal solution for 2 b hours. No treatment 1 2 3 k Isopropyl ester $ 2,i>-D, tr ie than ol amine s a lt 11 12 13 Ik 15 Nontreated 1 - 3 broad-leaved c a tta il k-% narrow-leaved c a tta il 6 7 8 9 10 2 -,1|-D, isopropyl ester 16 17 18 19 20 Treated 6 - 7 , 1 1 - 1 2 , 1 6 -1 7 narrow-leaved c a tta il 8 - 1 0 , 13-1 5 , 1 8 -2 0 broad-leaved c a tta il Plate I I . Five weeks a lte r c a tta il cut le a f tips were immersed in jars of commercial herbicidal solution for 2 k hours. No treatment 2 , 1*-D, isopropyl ester 2 6 3 k 8 10 ¥ & 2,i|-D, triethanolamlne sa lt 11 12 13 lh 1? 16 Nontreated 1 -2 narrow-leaved c a ttail 3-5 broad-leaved c a ttail 2,I|., 5-T, isopropyl ester 17 18 19 20 Treated 6 - 7 , 1 1 - 1 2 , lc -1 7 narrow-leaved cattail 8 - 1 0 , 1 3 - 1 5 , l 8—20 broad-leaved cattail Flate III'. Five weeks after c a tta il roots were immersed for 2k hours in 10 ppm of the phenoxyacetic acid formulation indicated,, i Narrow-leaved cattail Broad-leaved cattail Vegetative appearance Plate IV, , Narrow-leaved cattail -Broad-leaved cattail Fruiting bodies Vegetative and fruiting appearance of broad- and narrow-leaved cattail plants, Plate V. Portion of narrow-leaved c a tta il rhizome cross section {?00 X showing starch grains within c e lls. 34 Plate VX. C attail plants treated with systemic herbicidal formulations. 1056 1057 1006 1007 3 weeks a fte r treatment 1066 106U 1016 101U U weeks a fte r treatment Montreated 1006, 1 0 6 6 narrow— leaved c a t t a i l 1 0 0 6 3 1 0 1 6 broad— leaved c a t t a i l Tr eatment-— 8— 2U— 50 106U 1 2 .6 5 lb 2 ,U-D, triethanolamine s a lt 1 0 .5 6 lb 2 ,Lj— D, isopropyl ester 101U 1 5 .7 1 lb 2 ,Ll-D, triethanolamine s a lt 13.11 lb 2 ,b— D, isopropyl ester 1065 3.92 lb 2,U-D, Na s a l t 1015 1 U.3 2 lb 2 ,U-D, Na s a lt Treatment—'7— 7— 50 1051 25.30 lb 2 D, Na s a l t 1066 1065 1 01 6 1015 1007 J4I.I 46 lb 2jU— D, Na s a lt U weeks a fte r treatment Plate VII 0 1066 C a tta il plants treated -with systemic herbicidal formulation 1067 1016 1017 U weeks a fte r treatment 1066 1070 1016 1020 b weeks a f te r treatment Nontreated 1 0 6 6 narrow-leaved c a t t a i l 1 0 1 6 broad— leaved c a t t a i l Treatment-— 8— 2U— 50 1067 13*69 lb 2jl+-D, Na s a l t 1017 17.55 lb 2,h ~ D , Na s a l t 1070 712.00 lb pentachlorophenol 1 2 79.6 gal xylene 3.0. }_;9 gal C alifornia Spray Ortho No. 5 1020 571o93 lb pentachlorophenol 1027.9 gal xylene 1 0 . U9 gal California Spray Ortho No. 5 B Plate VIII. Treatment room and culture area following treatment with the various chemical formulations. A. Phenoxyac.etic acid treatment room B. Tank cultures follov/ing treatment v/ith phenoxyacetic acid and other chemical formulations Plate XX. Portion of stem or rhizome cross sections of troublesome ■western waterweeds. A. American pondweed rhizome 163 X B. Leafy pondweed stem. 330 X C. True waterweed stem l 6 f? X B c Plate X. Portion of stem or rhizome cross sections of troublesome ■western waterweeds . A. B. C. Slender sago pondweed stem l 6 £ X Horned pondweed stem 16£ X Richardson1s pondweed rhizome 165 X Plate XI. Systemic herbicidal treatment on water sedge. A. Water sedge (Carex aquatilis Wahl.) 5 weeks after 1st treatment B. 7 weeks after 2d treatment Treatment 1— ~6->23— (Left tank—front) 27.20 lb 2i h~Vs triethanolamine s a lt Treatment 2—8-L-50 Ii.5i| lb 2, 1-1.— D, triethanolamine s a lt 3.78 lb 2,lj-D, isopropyl ester Comments 9-22-50 2,Iu 5-T treated material Harvested Living % Dead % plant analysis 19.2 g 21.1 71.& g 78.9 Treatment 1—6-23-50 (Left tank—back) 30.29 lb 2,i|,5-T, isopropyl ester Treatment 2—8— U-59 1U.39 lb 2,li,5-T, isopropyl ester 2,[i-D, s a lt, ester treated material Living. % Bead T fg X lt W Tg W Z B Plate XIIo c Field photographs of three species of pondweedo A. Leafy pondweed (Potamoge ton foldosus Raf o) B* Slender sago pondweed (Potamogeton pectinatus L.) G. American pondweed (Potamoge ton nodosus Poiret) c Plate XIII* Systemic herbicidal treatment on leafy pondweed. Treatment 1 —S-.25-50 It-.93 lb 2 Jk-'E> Na s a l t Exposxire : 3 hr 35 min A. Leafy pondxveed (Potamogeton foliosus Raf•) Before treatment 8-2U-50 B. 1 week a fte r 1 st treatment C. 6 weeks a fte r 1 st treatment Comments 1 0-.6-50 Estimated survival. 0 % c Plate XIV. Systemic herbicidal treatment on- leafy pondweed„ Treatment 1— — G— 2|?— .65 Tb 2 y )4-D, isopropyl ester 6.77 lb 2 ,l4-D, triethanolamine s a lt Exposure: 3 hr h& rain A. Leafy pond-vveed (Potamogeton foliosus Raf.) Before treatment 0 — 2 I4— 50 B. 1 week a fte r 1 st -treatment C. 6 vreeks a fte r 1st -treatment Comments 10-»6-»^0 Estimated survival 5> % P l a t e XV. S y s t e m ic h e r b i c i d a l t r e a t m e n t on l e a f y p o n d w e e d . Treatment. 1 —8 — 25-5>0 3. U9 lb 2 , I i , 5 - T , " isopropyl ester Exposure; 6 hr 20 min A. Leafy pondweed (Potamogeton foliosus Raf•) Before treatinerrtTTj^2TT-5>0 Bo 1 week afte r 1 s t treatment C. 6 weeks a fte r 1st treatment Comments 10-6— 5>0 Estimated survival 85 Plate XVI. Vegetative regrowth after cutting leafy pondweed Treatment 1—7-7-50 Proximal half of plants cut off at ground line with scissors Exposure: 5 hr 30 min A. Leafy pondweed (Potamogeton foliosus Raf.) Before treatment 7-7-50 B. 1 week after 1st treatment C. b weeks after 1st treatment D. 7 weeks after 2d treatment Treatment 2-8-7-50 Plants in front half of tank cut off a t ground line with scissors Exposure: U hr 50 min Comments 9— 29-50 Estimated survival 100 % Fresh weight 2l(.8.0g Oven-dry weight 30.3 Percent oven-dry 12.2 weight C Plate XVII. D Systemic herbicidal treatment on American pondweed. Treatment 1—6-17-50 6 „Bl lb 2 , B-Dy triethanolamine s a lt Exposure: 3 hr I;5 min A. American pondweed (Potamogeton nodosus Poiret) Before treatment 6-15-50 B. Uweeks after 1st C. D. 6 weeks after 1st treatment 7weeks after 2d treatment treatment Treatment 2—8-lj.-50 6 .8 1 lb 2, B-Dy trie thanolamine s a lt Exposure: 3 hr 25 min Comments 9-25-50 Estimated survival 2 Cf A> Fresh weight Uo.5 Oven-dry weight 3.7 Percent oven-dry 9.1 weight Plate XVIII. C D Systemic herbicidal treatment on American pondweed, Treatment 1—6-17-5>0 538 lb 2>-D, isopropyl ester Exposure: 5 > lir 10 min A. American pondweed (Potamogeton nodosus Poiret) Before treatment &-l5~5>0 B. I;weeks after 1 s t treatment G. 6 -weeks after 1 s t treatment D. 7weeks after 2d treatment Treatment 2-— 8-L|.-5>Q 5*88 lb 2Jb-'Qs isopropyl ester Exposure : 3 hr 35 min Comments 9-25>-50 Estimated survival % % Plate XIX. c D Systemic herbicidal treatment on American pondweed, Treatment 1-— 6— 17— E>0 .9 . US lb 2 9\±9 5>-T* isopropyl ester Exposure: 3 hr 2£ min A. American pondweed (Potamogeton nodosus Poiret) Before treatment 6 — 15?— £0 B. k C. 6 weeks after 1st treatment D. 7weeks after 2d treatment v/eeksafter 1st treatment Treatment 2--8-U-50 7 . 3 2 lb 2 >U>5 ~T, isopropyl ester Exposure: 3 hr 15> min Comments 9-22-3>0 Estimated survival 5> % Fresh weight 53»7g Oven— dry weight 3o6 Percent oven-dry 6o7 weight C Plate XX. D Vegetative regrowth after cutting American pondweed Treatment I-.-7 .-7 ..5X ) Plants in front half of tank cut off at ground line with scissors Exposure t 5 hr U5 min A. American pondweed (Potamogeton nodosus Poiret) Before treatment 7-7-’50 B. 1 week after 1st treatment C. U weeks after 1st treatment D. 7 weeks after 2d treatment Treatment 2— .8-7— 50 Plants in front half of tank cut off at ground line with scissors Exposure t 6 hr 20 min Comments 9-29-50 Estimated survival 100 % Fresh weight 283.3g Oven-dry wei ght 30.2 Percent oven-dry 10.7 weight Plate XXI. Systemic herbicidal treatment on gigantic sago pondweed, Treatment 1 —6-17-5>0 6.91 lb 2,'ii-D, triethanol amine s a lt Exposure: h hr 20 min A. Gigantic sago pondweed (Potamogeton pectinatus Lo) Bexore treatment 6-15-50 B. h weeks after 1st treatment C. 6 weeks after 1st treatment D. 7weeks after 2d treatment Treatment 2—8-U-50 lb trie thanolamine sa lt Exposure: 3 hr 10 min Comments 9-22-^0 Estimated survival 2 % c Plate XXII. D Systemic herbicidal treatment on gigantic sago pondvreed. Treatment 1—6-17-50 Treatment 2—8-U-50 JcB'B l b 2 f h-D} isopropyl ester k *97 lb 29 U^Df isopropyl ester Exposure: Exposure: 3 hr 35 min Comments 9-22-50 Estimated survival 1 % 5 hr lj.0 min A. Gigantic sagopondweed (Potamogeton pectinatus L.) Before treatment 6-l5-5o B. h "weeks after 1st treatment C. 6 weeks after 1st treatment D. 7weeks after 2d treatment Plate XXXII. Systemic herbicidal treatment on gigantic sago pondweed. Treatment 1—9-15-50 10."62 ib ' 2 / U j i s o p r o p y l ester Exposure: 3 hr 50 min A. Gigantic sago pondweed (Potamogeton pectinatus L .) Before treatment 9-l5~5>0 B. 3 weeks af'ber 1st treatment C. 5 weeks after 1st treatment Comments 10-20-50 20 Estimated survival 5U.2g Fresh weight 3.2 Oven-dry weight Percent oven-dry 5.9 weight c Plate XXIV. 33 Vegetative regrowth a fte r cutting gigantic sago pondweed. Treatment 1~-.7-lU-^O Plants in front h alf of tahlc cut off a t ground lin e with scissors Exposure: 5 hr 25 min A. Gigantic sa.go pondweed (Potamogeton pectinatus L .) Before treatment 7— lU-5’0 Bo 2weeks a fte r 1 s t treatment C, 6 weeks a fte r 1 st treatment D. Uweeks a fte r 2d treatment Treatment 2-.-.8-22~5>0 Plants in front h alf of tank cut off a t ground line with scissors Exposure: h hr min Comments 9-25>— 5>0 Estimated survival 100 % Plate XXV. Contact herbicidal treatment on slender sago pondweed, Treatment 1—7-7-50 Zf+sT.i+O lb NaAs02 Exposure: U hr 35 min A. Slender sago pondweed (Potamogeton pectinatus L.) Before treatment 7-7-50 B. 7 weeks after 2d treatment Treatment 2-8-7-50 27i-u7U lb NaAs02 Exposure: 3 hr 35 min Comments 9-29-50 Estima'ted survival 25 % Fresh weight 9«2g Oven-dry weight 0.5 Percent oven-dry 5«U weight Plate XXVI. C Systemic herbicidal treatment on horned pondweed. Treatment 1—S-.25-l?0 S'.6 £ lb 2VH-DV"isopropyl ester 6 . 7 7 lb 2 ,U-Da triethanolamine s a lt Exposure: 3 hr 20 min A. Horned pondweed (Zannichellia p alu stris L .) Before treatment 6-2ir-^0 Bo 1 week after 1 s t treatment C. 6 weeks after 1st treatment Comments IO-6-5G Estimated survival 0 % Plate XXVIi. Systemic herbicidal treatment on horned pondweed. Treatment 1 — .8-25-50 7*20 lb 2 s i\.i S - r^y isopropyl ester Exposure: 5 hr 20 min A. Horned pondvreed (Zannichellia palustris Lo) Before treatment 6 — 2a— 3>0 B. 1 week after 1st treatment G. 6 weeks after 1st treatment Comments 10-6-50 Estimated survival