The influence of Nitrogen on seedling and early growth of perennial ryegrass and Cocksfoot

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New ealand Jurnal f Agricultural Research SSN: 0028-8233

O'Brien (1960) The influence f Nitrgen n seedling and early

Agricultural Research, 3:2, 399-411, DO: 10.1080/00288233.1960.

1960.10418095 Published nline: 21 Dec 2011.

1 New ealand Jurnal f Agricultural Research SSN: (Print) (Online) Jurnal hmepage: The influence f Nitrgen n seedling and early grwth f perennial ryegrass and Ccksft T. A. O'Brien T cite this article: T. A. O'Brien (1960) The influence f Nitrgen n seedling and early grwth f perennial ryegrass and Ccksft, New ealand Jurnal f Agricultural Research, 3:2, , DO: / T link t this article: Published nline: 21 Dec Submit yur article t this jurnal Article views: 146 Citing articles: 3 View citing articles Full Terms & Cnditins f access and use can be fund at

2 1960) 399 THE NFLUENCE OF NTROGEN ON AND EARLY GROWTH OF PERENNAL AND COCKS FOOT SEEDLNG RYEGRASS By T. A. O'BREN, Grasslands Divisin, Department f Scientific and ndustrial Research, Palmers tn Nrth (Received fr publicatin, 11 January 1960) Summary The grwth f seedlings f perennial ryegrass (Llium perenne L.) and ccksft (Dactylis glmerata L.) was studied at different sil nitrgen levels. At all nitrgen levels, and at all stages f grwth studied, perennial rye grass seedlings prduced mre dry matter than ccksft. After the initial tw-weeks perid, the dry-matter prductin rati f perennial rye grass t ccksft was greater at the lwer levels f nitrgen than at the higher. At high nitrgen levels the relative grwth rates f the tw species were similar, but at lwer nitrgen levels the relative grwth rate f perennial ryegrass was significantly greater than that f ccksft in the secnd tw weeks f grwth, and significantly less than that f ccksft in the furth tw weeks f grwth. The results are discussed in relatin t early cmpetitin as affected by sil nitrgen, light, and temperature, between these tw species in pasture mixtures. NTRODUCTON n New ealand, attempts t establish mixtures f perennial ryegrass (Llium perenne L.) and ccksft (Dactylis glmerata L.) in sils f high fertility, are ften unsuccessful because the ryegrass suppresses the ccksft. Where available nitrgen is at intermediate r lwer levels perennial ryegrass seedlings appear t cmpete less strngly with thse f ccksft. This difference appears t be greater with autumn swings than with spring swings. Pasture species have been classified by varius wrkers accrding t their seedling aggressiveness. Stapledn and Davies (1927) and Blaser et al. (1956), n the criterin f relative weights f seedlings at sme time after swing and grwn under cnditins f high fertility, rated perennial ryegrass as being mre aggressive than ccksft. Factrs discussed by these wrkers as cntributing t this seedling aggressiveness included seed size, rate f germinatin and rate f seedling grwth. Cashmre (1934) investigated sme factrs f the aggressiveness f perennial ryegrass ver Phalaris tubersa L. in high fertility sil in the first grwing seasn. Using bth single plants and mixtures in plts, he cncluded that the depressing effect f ryegrass n phalaris in the seedling year was due t the greater tp grwth f the frmer cmbined with its denser rting system in the upper ft f sil. N.. J. agric. Res. 3:

400 NEw EALAND JOURNAL OF AGRCULTURAL RESEARCH (APRL Cmpetitin fr light and nutrients under cnditins f liberal water supply have been demnstrated by several wrkers t be the cntributing factrs t a

3 400 NEw EALAND JOURNAL OF AGRCULTURAL RESEARCH (APRL Cmpetitin fr light and nutrients under cnditins f liberal water supply have been demnstrated by several wrkers t be the cntributing factrs t a species aggressiveness during seedling grwth. Chippindale ( 1932) attributed the aggressiveness f talian ryegrass (Lalium multiflarum L.) ver meadw fescue (F estuca pratensis L.) t this, and Dnald (1958) cncluded that the same factrs (the nutrient in this case being nitrgen) enabled perennial ryegrass t be the aggressr species ver phalaris. Furthermre, Dnald shwed that the cmbined effect f cmpetitin fr bth factrs exceeded the sum f the separate effects. This present study was undertaken t determine the influence f sil nitrgen level n the cmparative grwth f perennial rye grass and ccksft at varying stages f their seedling and early grwth. EXPERMENTAL The experiment was carried ut at the Palmers tn Nrth statin f Grasslands Divisin, Department f Scientific and ndustrial Research. Fig..-One blck f the layut, shwing 10 randmised plts and tw sub-plts per plt fr the tw species. The temperature recrding unit is als shwn.

4 1960) O'BREN-NFLUENCE OF NTROGEN 401 The experimental site was divided int six blcks each measuring 21 ft x 3 ft (see Fig. 1). Each blck was subdivided int ten plts t prvide a ttal f 60 plts, each measuring 2 ft x 3 ft. The sil used fr all plts was Taup pumice alluvium (Fleming 1953). This sil type (ttal nitrgen 0.02%) was chsen fllwing a preliminary glasshuse experiment designed t determine the degree f plant respnse t nitrgen treatment in different sil types. Each plt was divided int tw sub-plts ne f which cntained tw rws f perennial ryegrass and the ther tw rws f ccksft, allwing six inches between all rws within the plt. Sixteen unifrmly germinated seeds f bth species were swn per rw at tw-inch spacings and at i in. depth, n 21 Nvember T btain cmmn emergence time and physilgical age at transplanting, it was necessary, frm the results btained in a preliminary investigatin n the differences in rate f germinatin between these tw species, t pre-germinate the ccksft seed tw days befre the ryegrass. This was dne n sand in petri dishes (in half-strength Hagland's slutin) and seed was then selected fr unifrmity, at the stage f emergence f the clerhiza. The term "germinated seeds" referred t hereafter dentes this stage, when actual planting tk place. Dry weights f germinated seeds were determined by drying and weighing ten grups (each f ne hundred) f unifrmly germinated seeds fr each species. Nitrgen treatments applied were as fllws:- 1 1 CONTROL 6 b N/ac. 20 lb N/ac. 8 S 9 60 b N/ac b N/ac. Nitrgen was applied in the frm f ammnium nitrate (35% N). These were laid dwn as a 10 X 2 split-plt design, the additinal replicates within blcks being used t prvide greater accuracy in measurement at the lwer levels f N. The numbers f plants per harvest fr each treatment were therefre: 144 fr Cntrl and 6 lb N/ac, 96 fr 20 lb N/ac, and 48 fr 60 and 120 lb N/ac. Superphsphate at 5 cwt/ac, and magnesium sulphate and muriate f ptash at 1 cwt/ac, were applied t all plts. The superphsphate was applied five days prir t swing and the quantity t be given per plt was bulked with pumice and spread evenly, at apprximately i in. depth alng the drill rws. Bth the muriate f ptash and magnesium sulphate were applied in slutin tw days prir t swing.

402 NEw EALAND JOURNAL OF AGRCULTURAL RESEARCH (APRL Nitrgen was applied in split applicatins in the fllwing way. One tenth f the whle f each treatment was added ne day prir t swing.

5 402 NEw EALAND JOURNAL OF AGRCULTURAL RESEARCH (APRL Nitrgen was applied in split applicatins in the fllwing way. One tenth f the whle f each treatment was added ne day prir t swing. One day subsequent t seedling emergence fr bth species, ne twentieth f the whle f each treatment was applied in slutin and additins at this rate were repeated with such frequency and at regular time intervals, that by the end f the third wek all nitrgen treatments had received half f their prpsed applicatin. The balance f the applicatins were made twice weekly, again at the rate f ne twentieth f the whle treatment, fr the remainder f the experimental perid, the last applicatin being given fur days prir t the final harvest. All plts were watered at frequent intervals thrughut the experimental perid. Fur harvests were made at frtnightly intervals, the first being made furteen days after mean seedling emergence. There was a delay f apprximately ne day in ccksft emergence, but this has been ignred. At each harvest date, eight plants were harvested frm each sub-plt, fur being taken frm each rw, and bulked. These plants were selected frm a restricted randm plan in such a way that after the secnd harvest the distance between the plants was dubled. At each harvest, the plants frm each sub-plt were washed, rts trimmed ff, and the remainder bulked in air-tight cntainers and put in cl strage. After determinatin f tiller numbers, leaf numbers, and leaf areas, the shts were ven-dried and weighed. The grwth increments between harvests were determined, the seed weights being used t determine the increments at the first harvest. The results were then expressed in terms f relative grwth ra tes (Blackman 1919). The methd f determining leaf area was as fllws. Twenty plants f each species frm each treatment were selected at randm at each harvest. All leaves frm each plant were severed at the psitin f the ligule, and frm these a number f bth mature and immature leaves were selected. At the middle f each selected leaf, a 2 cm sectin was cut and its width measured, the rest f the leaf being bulked with the remaining leaves fr each plant. Bth the bulk f the leaf material, the strips and the stems were ven-dried and separately weighed. The ttal leaf area fr each plant was then calculated. Weather Cnditins RESULTS Mid Nvember t mid January, a perid with apprximately a 16-hur day length, was n the whle fine and sunny. The means f the daily maximum and minimum temperatures (OF) fr the perid were:- 6 in. abve Grund Surface in. belw 3 in. belw Maximum Minimum

6 TABLE 1. DRY WEGHTS (mg) OF THE SHOOTS OF SNGLE PERENNAL RYE GRASS AND COCKSFOOT PLANTS AS NFLUENCED BY NTROGEN LEVELS AT FOUR HARVEST DATES. RYEGRASS/COCKSFOOT RATOS ARE SHOWN N TALCS Treatment (Equivalents Harvest 1: 9/12/58 Harvest 2: 23/12/58 Harvest 3: 6/1/59 Harvest 4: 20/1/59 f Applied P. P.. P' Nitrgen in erenma Ccksft erenmal Ccksft Perenmal! Ccksft, erenma lb N per acre) Ryegrass Ryegrass Ryegrass i Ryegrass Ccksft Nil 2.48 ± 0.06 j 1.20 ± 0.03) 14.4 ± ± ± ± ± ± " Cefficien t f variatin (per 8 plants) / ) % 4.5% 4.6% 49 ] %... (,0 0'> t'1... "1 :" c:: t'1 t'1 "1 ::i g t'1 >> (,,)0

7 404 NEW EALAND JOURNAL OF AGRCULTURAL RESEARCH (APRL These recrdings were btained frm a recrding unit sited in the plts (see Fig. 1). The maximum temperature recrded, which was at the sil surface, was F n 11 January. During the same day the maxima recrded fr six inches abve the sil surface, ne inch belw and three inches belw were 90.5 F, 86 F and 82.5 F respectively. The minimum sil temperature recrded was 33 F n 21 Nvember. The minima at the sil surface, ne inch and three inches belw the sil surface during this same day were 34 F, 38 F and 48 F respectively. Seed Weight's The ttal dry weights (g) f ne thusand germinated seeds, tgether with standard errrs, were:- Perennial rye grass Ccksft Herbage Yields (Dry Matter) Fr each harvest, cefficients f variatin were stable ver the five treatments. The data analysed are therefre lgarithms f mean weights (mg) per plant. The interpretatins were made frm the analysis f variance and transfrmed means, but in the table f means presented in Table 1 the decded means and standard errrs are given. The fllwing are the significant features f Table 1: (a) Perennial ryegrass ut yielded ccksft in all treatments at all harvest dates. (b) There were prductin increases frm added nitrgen fr bth species at all harvests. (c) There were significant species x nitrgen level interactin> at all harvests, i.e., the yield rati f ryegrass t ccksft changed significantly with nitrgen level at all harvests. Table 1 als shws that the abslute difference in favur f perennial ryegrass increased prgressively with nitrgen level with n sign that the difference between the species had reached its maximum at 120 lb N/ac. Tiller Number Tiller numbers fr the fur harvest dates are presented graphically (lg. scale) in Fig. 2. Average S.E.s are als shwn fr each harvest. The fllwing cmparisns are f interest: (a) At all harvest dates, and at all nitrgen levels, there were species differences in tiller number in favur f perennial ryegrass. The nly exceptin was at the cntrl level in the first harvest, in which the difference did nt reach significance. (b) At all harvests, there were significant species x nitrgen interactins, the abslute increase in tiller number with increased nitrgen being significantly greater in perennial ryegrass.

8 1960) O'BREN-NFLUENCE OF NTROGEN 405 : 50' '0 25' 0 cr W CD ::;: 10-C :J cr 5'0 w -'.-- _ <1..J ::;: 1 0 O 51 i i C'l ' 0 14 DAiS FROM EMERGENCE 56 S.E. 8A 0'24 0'14 Fig. 2.-Legend: Perennial ryegrass, ; Ccksft, Be. Numbers f tillers prduced by single perennial ryegrass and ccksft plants in respnse t nitrgen levels, at fur harvest dates. Leaf Number Leaf numbers fr the fur harvest dates are shwn graphically in Fig. 3 (lg. scale). Average S.E.s are als shwn. The cnclusins t be drawn are the same as thse given abve fr tiller number.

9 TABLE 2. LEAF AREAS (sq. em) OF SNGLE PERE\:-<A. RVEGRASS A:-<D COCKS FOOT PLANTS AT THE FOUR HARVEST DATES (r;:l:s Harvest 1: 9/12/58 i_vst 2/12/58 Harvest 3: 6/1/59 Harvest 4: 20/1/59 f Applied P' P. P' P' Nitrgen in erenma Ceksft erenma Ccksft erenmal Ccksft erenma Ceksft b N per acre) Ryegrass Ryegrass Ryegrass Ryegrass Nil i Ol i:'l N i:'l :> t...; c:: '" z :> t"' "l > 6 c:: t"' >j c:: t"' i:'l UJ i:'l >- '" "" p

10 1960) O'BREN-NFLUENCE OF NTROGEN a:: w! :1: ::> z / / / / / SA Be Be / -Be LL <l: W...J «t.j DAYS 14 FROM EMERGENCE 56 S.E O g Fig. 3.-Legend: Perennial rye grass, ; Ccksft,. Numbers f leaves prduced by single perennial rye grass and ceksft plants in respnse t nitrgen levels, at fur harvest dates. Leaf Area Mean leaf areas per plant are presented in Table 2. The fllwing pints are f interest: (a) The increase f leaf area in perennial ryegrass in resp:m')c t increasing levels f nitrgen appeared t be twice that f ccksft fr the first three harvest dates, except at 20 lb N / ac in the third harvest when the increases were similar in the tw species.

11 408 NEW EALAND JOURNAL OF AGRCULTURAL RESEARCH (APRL (b) At the furth harvest, the differences in leaf area between the species had narrwed appreciably except at the highest nitrgen level, where the difference cntinued t favur perennial ryegrass. Relative Grwth Rate Relative grwth rates, tgether with their standard errrs, are s!lown graphically in Fig '20 > < Q...,. 0'1' "...,. "e '" 0 0 1" W 0"2 i 0 10 " e :! 0'02 PERENN AL RYEGRASS COC<SFOOT _0_0_ : i _ /! / / T / // ( (/ t EQUVALENTS OF APPLED NTROGEN (LB./ ACRE) 0-' WEEKS 0'22 > 0 20 < "J 0'U5 0". 0'12 : i /1/: j 0 08 t 0 0 0(5. PERENNAL 0 0" RYEGRASS 11_ -11'-- W 0'02 COCKSFOOT _0_0_ ' EurVA.LENTS OF.4PPLED NTROGEN (Le./ ACRE) 2-' WEEKS 0'20 > ;l... 0'18 ': PERENNAL RYEGRASS - -)1---)1-- COCK$FOOT -0--' Q "4 PEAENNA.L RYEGRA5S - -)1- - _t COCKsrOOT W :. 0"2 0 0 '0 ;: 0'08 " z DlO i 0'08 " 0'01S w > g 0'0" 0'02 e EQUVALENTS OF' APPLED NTROGEN (L&./.A.CRE) "-8 WEEKS EQUVALENTS r APPLED NTROGEN (Le./ ACRE) 15-8 WEEKS Fig. 4.-Relative grwth rates during the fur intervals between harvests (mg/mg/day) f perennial ryegrass and ccksft as influenced by nitrs-en levels.

12 1960) O'BREN-NFLUENCE OF NTROGEN 409 The fllwing are significant at the 5% level. (a) At the first harvest, in all except the highest level f nitrgen, ccksft was shwn t have higher relative grwth rates than perennial ryegrass. At the highest level f nitrgen, there was n real difference between their relative grwth rates. t shuld be nted that dry weights f germinated seeds were used as base lines fr calculatin f these relative grwth rates. These seed weights may nt represent plant embrys adequately in either species, and may even intrduce a bias int species cmparisns. Cnclusins fr this harvest shuld therefre be treated with cautin. (b) At the secnd harvest, at the lwer nitrgen levels, perennial ryegrass shwed higher relative grwth rates than thse f ccksft. At the tw higher nitrgen levels there were n real differences. (c) At the third harvest, at all nitrgen levels except the cntrl, ccksft was shwn t have higher relative grwth rates than thse f perennial ryegrass. (d) At the furth harvest, ccksft had higher relative grwth rates than thse f perennial ryegrass at the lwer levels f nitrgen, but at he highest level f nitrgen the difference did nt reach significance. DSCUSSON This experiment was carried ut during early summer, a time f the year when prevailing temperatures favur the grwth f ccksft (Mitchell 1956), and the amunt f incident light energy available fr grwth is at a maximum. Mrever, because f the watering prcedure used during the experiment, sil misture levels were at all times high, and adequate fr maximum grwth. Under these climatic cnditins the relative perfrmance f the seedlings and early grwth f ccksft cmpared with perennial ryegrass wuld be as high as culd be apprached under field cnditins. Hwever, even under the experimental cnditins, perennial ryegrass shwed greater seedling aggressiveness than ccksft when judged by plant size and plant yield at all nitrgen levels studied. Table 1 shws that, at all stages f grwth and all nitrgen levels, perennial ryegrass prduced mre than twice as much weight f sht tissue per plant as ccksft, the difference in favur f perennial ryegrass increasing prgressively with nitrgen level. There was ne exceptin t this where ryegrass yielded nly ne and a half times as much as ccksft. Similarly, rate f tiller prductin and increases in leaf number in perennial ryegrass shwed the same trend with respect t nitrgen level. Als increases in leaf area in perennial ryegrass were twice thse f ccksft fr the first three harvests, althugh, at the furth harvest, the differences in leaf area between the species narrwed appreciably, except at the highest nitrgen level, where the differences cntinued in favur f perennial ryegrass.

410 NEW EALAND JOURNAL OF AGRCULTURAL RESEARCH (APRL t is cnsidered that the significant difference in rapidity f tiller and leaf prductin between the tw species is a genetic difference and is due,

13 410 NEW EALAND JOURNAL OF AGRCULTURAL RESEARCH (APRL t is cnsidered that the significant difference in rapidity f tiller and leaf prductin between the tw species is a genetic difference and is due, as suggested by Sharman (1942, 1947) t differences between the ptential numbers f leaf and tiller primrdia capable f being prduced in these tw species. Als the cntrast in leaf area develpment between treatments at the same stage f grwth was induced, it may be reasnably assumed, by the large differences in the prtein nutritinal status f the plants. This is in accrd with Wardlaw's view (1952) that the dimensins f the leaf are strngly influenced by the nutritinal status and hence activity (Mitchell and Sper 1958) and size (Wardlaw 1952) f the sub-apical meristematic tissue. When the abve results are cnsidered in relatin t field practice, they shw that perennial ryegrass is a mre aggressive species in the seedling and early vegetative stage f grwth than ccksft, even in sils f very lw fertility and under cnditins where climatic factrs favur maximum grwth f ccksft. At high nitrgen levels, cmparable with sils f high fertility, and at all ther seasns f the year when cnditins are less favurable fr ccksft, the aggressiveness t perennial ryegrass wuld be mre prnunced. Frm the results it wuld appear that if ccksft is t be successfully established, with perennial ryegrass, then lw seeding rates, with ccksft as the majr grass cmpnent in the seeds mixture shuld be emplyed regardless f sil nitrgen levels. f plants f equal size are cmpared, the cmpetitive advantage changed in favur f the ccksft after the furth week f grwth at the lwer nitrgen levels, wing t the fact that ccksft after this time had a higher relative grwth rate than perennial ryegrass (Fig. 4). Frm the abve result, it wuld appear that a successful methd fr achieving a balanced ryegrass/ccksft/clver pasture n sil f high r medium fertility, wuld be first t lwer the sil fertility by crpping. The mixed grass/clver pasture culd then be established by verswing the ryegrass at a slightly later stage int an already develping ccksft/clver sward, s that plants f apprximately equal size are btained when the first grazing is desired. The sequence f verswing the ryegrass int a ccksft/clver sward wuld als appear t be f added imprtance in establishing a new pasture n a sil lw in available nitrgen, as certain grass species by virtue f their ready germinatin and early rapid grwth in sils lw in available nitrgen are able t cmpete mre vigrusly fr nitrgen and at an early stage exhaust the limited sil nitrgen supply thereby suppressing the grwth f assciated species (Sears 1953; Dnald 1958). Table 1 shws that ccksft wuld be less cmpetitive tward clvers fr nitrgen than ryegrass, at the stage when legumes are dependent upn sil nitrgen fr grwth befre they becme independent thrugh symbitic nitrgen fixatin (Wilsn 1940). ACKNOWLEDGMENTS The authr is indebted t Dr P. D. Sears, Directr, Grasslands Divisin, fr suggesting the prject and arranging facilities fr the study,

14 1960) O'BREN-NFLUENCE OF NTROGEN 411 t the late Mr E. O. C. Hyde and Mr R. W. Brugham fr help and guidance, and t Miss M. A. Leavey fr technical assistance. The assistance f Mr A. C. Glenday, Bimetrician, Applied Mathematics Labratry, Department f Scientific and ndustrial Research in statistical analyses is much appreciated. REFERENCES Blackman, V. H. 1919: The Cmpund nterest Law and Plant Crwth. Ann. Bt. Lnd. 33: Blaser, R. E.; Taylr, T.; Griffeth, W.; Skrdla, W. 1956: Seedling Cmpetitin in Establishing Frage Plants. ]. Amer. Sc. Agrn. 28: 1-6. Cashmre, A. B. 1934: A Cmparative Study f Llium perenne L. and Phalaris tubersa L. at Varying Stages f Grwth. Cmmnw. sci.industr. Org. (Aust.) Bull. 81. Chippindale, H. G. 1932: The Operatin f nterspecific Cmpetitin in Causing Delayed Grwth f Grasses. Ann. appl. Bii. 19: Dnald, C. M. 1958: The nteractin f Cmpetitin fr Light and fr Nutrients. Aust.]. agric. Res. 9: Fleming, C. A. 1953: The Gelgy f Wanganui Sub-divisin. N.z. gel. Surv. Bull. n.s. 52. Mitchell, K. J. 1956: Grwth f Pasture Species under Cntrlled Envirnment.. Grwth at Varius Levels f Cnstant Temperature. N..]. Sci. Tech. A38: Mitchell, K. J.; Sper, K. 1957: Effects f Differences f Light ntensity and Temperature n the Anatmy and Develpment f Leaves f Llium perenne L. and Paspalum dilatatum Pir. N.. ]. agric. Res. 1: Sears, P. D. 1953: Results f Pasture Establishment Trials. Prc. 15th Cnt. N.z. Grassl. Ass Sharman, B. C. 1942: Sht Apex in Grasses and Cereals. Nature, Lnd. 149: : Bilgy and Develpmental Mrphlgy f the Sht Apex in the Gramineae. New Phytl. 46: Stapledn, R. G.; Davies, W. 1927: Seeds Mixture Prblems. Welsh Plant Breed. Sta. Bull. Ser. H.6. Wardlaw, C. W. 1952: Experimental and Analytical Studies f Pteridphytes. XV. The Nutritinal Status f the Apex and Mrphgenesis. Ann. Bt. Lnd. n.s. 16: Wils:m, P. W. 1940: "The Bichemistry f Nitrgen Fixatin". Univ. f Wiscnsin Press, 302 pp.

M.J. Guthriel, D.G. Wagner2

EFFECT OF SUPPLEMENTAL PROTEIN OR GRAIN ON INTAKE AND UTILIZATION OF MEDIUM QUALITY PRAIRIE HAY BY STEERS M.J. Guthriel, D.G. Wagner2 and D.C. Weakley3 Stry in Brief Sixteen 1,053 lb tw-year-ld Herefrd