Clonal Growth of White Clover: Factors Influencing the Viability of Axillary Buds and the Outgrowth of a Viable Bud to Form a Branch

The development of an axillary bud of white clover to form abranch depends on (1) the bud being viable, vegetative and non-dormant,and (2) suitable conditions for outgrowth of the bud. Foragingtheory emphasises the second of these requirements. Glasshousestudies with white clover rarely result in a loss of bud viability.In contrast, in field populations over 50% of the buds reachingthe stage of maturity when branching can occur are not in aviable, vegetative, non-dormant condition. We examined whethernon-viability could be induced in a glasshouse experiment byapplying treatments in a factorial design. The factors were:defoliation, phosphorus supply, soil moisture status, simulatedtreading and grass competition. In addition, we measured theeffects of the treatments on the outgrowth of viable buds inorder to assess whether the same factors were determining viabilityand outgrowth. Defoliation significantly reduced bud viability(by 44%) but no other factors, either singly or in combination,had a significant effect. A greater variety of factors and combinationsof factors influenced bud outgrowth; these were defoliation,phosphorus status and interactions involving phosphorus andgrass; defoliation, phosphorus and soil moisture; and soil moisture,grass and treading. For white clover it is relevant to includethe state of the axillary meristem in any model of foraging. Trifolium repens ; white clover; axillary bud; viability; clonal growth; foraging; defoliation     

Effect of Light Quality (Red:Far-red Ratio) at the Apical Bud of the Main Stolon on Morphogenesis of Trifolium repens L.

A controlled environment experiment investigated whether thered:far-red (R:FR) ratio of light at the apical bud of the mainstolon could alter plant morphogenesis in clonal cuttings ofwhite clover (Trifolium repens L.) The apical bud included theapical meristem, five to six developing leaf primordia withassociated axillary bud primordia and stipules and the firstemerged folded leaf until development was greater than 0·3on the Carlson scale. Three light regimes were imposed on theapical bud by collimating light from R or FR light-emittingdiodes so that the R:FR ratio of light incident at the apicalbud was set at 0·25, 1·6 or 2·1, withoutsignificantly altering photosynthetically active radiation.The effect of these light regimes on white clover seedling growthwas also tested. At a low R:FR ratio seedling hypocotyl and cotyledon lengthswere significantly longer. However, with the cuttings, the lighttreatments did not alter node appearance rate or internode lengthof the main stolon, petiole length, area of leaves or totalshoot dry matter. There was one significant photomorphogeneticresponse in the cuttings, a delay of 0·5 of a phyllochronin the appearance of branches from axillary buds in the lowR:FR ratio treatment relative to the other treatments. Wherebranch appearance was delayed plants had fewer branches. Thisdifference could be ascribed solely to a delay in branch appearanceas there were no significant treatment effects on either theinitiation of axillary bud primordia within the apical bud,the probability of branching or on the rate of growth of branchesafter appearance. Because treatment of the apical bud inducedonly one of the many previously observed responses of whiteclover to a decrease in the R:FR ratio of light, we concludethat other plant organs must also sense the quality of incidentlight.Copyright 1994, 1999 Academic Press White clover, Trifolium repens, apical bud, light quality, red:far-red ratio, light-emitting diode, branching, axillary buds, photomorphogenesis     

Development of Axillary and Leaf-opposed Buds in Rattan Palms

Axillary vegetative buds are present in Calamus, Ceratolobus,and Plectocomiopsis. Two species of Daemonorops Sect. Piptospathaalso have axillary vegetative buds. All species of Daemonoropshave only displaced adnate axillary inflorescence buds. A singlebud is initiated in the axil of the first or second leaf primordiumin a way similar to that for axillary inflorescence buds. Themeristem is displaced during development on to the internodeabove and sometimes on to the base of the leaf above. Leaf-opposedvegetative buds occur in five species of Daemonorops Sect. Cymbospathaand in one species of Daemonorops Sect. Piptospatha. This typeof bud is initiated 180° away from the axil of the firstor second leaf primordium. It is not a displaced axillary bud,but does become adnate to the internode above like the axillarybuds. One or more leaves, transitional between juvenile andadult, on a shoot often subtend both types of buds. Myrialepishas leaf-opposed vegetative buds, but their development wasnot observed. Korthalsia has buds that are displaced about 130°from the leaf axil and are intermediate between the axillaryand the leaf-opposed condition. Other forms of vegetative budsare described: multiple buds in Plectocomia, aerial forkingin Korthalsia, and suckering from inflorescences and from aerialstems in Calamus. bud development, rattan palms, palm taxonomy, branching     

Influence of Position and Number of Nodal Roots on Outgrowth of Axillary Buds and Development of Branches in Trifolium repens (L.)

The implications of the presence of a root, either at the parentnode or at neighbour nodes, on branch formation of Trifoliumrepens (white clover) was investigated. Plants were freely rootedor rooting was restricted to every sixth or every twelfth nodealong the parent axis. The absence of a root at the parent nodehad little influence on the probability of the subtending axillarybud forming a branch but, on average, delayed the outgrowthof the bud. The probability that an axillary bud, emerging froma non-rooted parent node, developed to a lateral branch (branchwith elongated internodes) decreased with decreasing proximityof the parent node to a rooted node. Lateral branches emergingfrom non-rooted parent nodes which were two nodes distal toa rooted node had a higher rate of node appearance, a greatermean internode length and area per leaf, and were more branchedthan lateral branches emerging from other non-rooted parentnodes. The dry mass of each single root and of branches grownat rooted parent nodes were significantly higher in plants withrestricted rooting than in freely rooted plants. Restrictionin the number of rooted nodes per plant increased the numberof inflorescences. It is concluded that the whole plant responseto restricted root formation was continuous growth of the parentaxis and compensatory growth of the branch at the rooted node.In general, growth was slow for axillary buds whose developmentwas dependent on the basipetal movement or cross-transport withinthe stolons of resources exported from roots. Trifolium repens (L.); white clover; axillary bud outgrowth; branch development; clonal growth; nodal root     

The Effect of Defoliation on the Nitrogen Economy of White Clover: Regrowth and the Remobilization of Plant Organic Nitrogen

Single plants of white clover (Trifolium repens) were establishedfrom stolon cuttings rooted in acid-washed silver sand. Allplants were inoculated with Rhizobium trifolii, and receivednutrient solution containing 0·5 mg ¹⁵N as either ammoniumor nitrate weekly for 12 weeks (i.e. 6 mg ¹⁵N in total). Plantswere then leniently defoliated or left intact, and the labelledN supply was replaced with unlabelled N. Lenient defoliationremoved fully expanded leaves only, leaving immature leaveswhich accounted for 50–55% of the total; growing pointnumbers were not reduced. Nodules, leaves and growing pointswere counted over the following 21 d period, and d. wts, N contents,and ¹⁵N enrichments of individual plant organs were determined. Defoliated plants had fewer nodules, but numbers of growingpoints were unaffected by defoliation. The rates of both leafemergence and expansion were accelerated in defoliated plants;in consequence the number of young leaves remained less thanin intact plants until day 21. Total dry matter (DM) and N accumulationwere less in defoliated plants, and a greater proportion oftotal plant DM was invested in roots. About 97 % of plant totalN was derived from fixed atmospheric N, but there was incompletemixing of fixed and mineral N within the plant. Relatively moremineral N was incorporated into roots, whereas there was relativelymore fixed N in nodules. There was isotopic evidence that Nwas remobilized from root and stolon tissue for leaf regrowthafter defoliation; approximately 2 % of plant N turned overdaily in the 7-d period after defoliation, and this contributedabout 50% of the N increment in leaf tissue. White clover, Trifolium repens L. cv. SI84, lenient defoliation, N economy, regrowth, N remobilization     

Correlative Inhibition in the Shoot of Agropyron repens ( L.) Beauv

Correlative inhibition was investigated in plants of Agropyronrepens at two temperatures. Reciprocal inhibition ocrurred betweenthe main shoot apex and the outgrowing axillary shoots, withthe balance of inhibition varying with temperature. Apical dominancewas stronger at 10 °C than at 20 °C , but even at 10°C release of apical dominance by decapitation had onlyminor effects on the timing of outgrowth, growth pattern andrate of dry weight aocumulation of the axillary shoots. Dominanceof the main shoot apex by the axillary shoots was stronger at20 °C than at 10 °C. Removal of axillary buds preventeddecline in size and activity of the main shoot apex ard resultedin increased rates of primordium initiation, leaf emergenceand dry weight accumulation in the main shoot. It is suggestedthat a system of reciprocal dominance provides a mechanism formaintaining the characteristic habit of the grass plant andlimits growth in height of vegetative shoots. Agropyron repens (L.) Beauv, couch grass, correlative inhibition, apical dominance, shoot, apex     

Growth of White Clover, Dependent on N2 Fixation, in Elevated CO2 and Temperature

Clonal plants of white clover (Trifolium repens L ), whollydependent on N₂ fixation, were grown for 6 weeks in controlledenvironments providing either (C680 regime) 23/18 °C day/nighttemperatures and a CO₂, concentration of 680 µmol mol^–1,or (C340 regime) 20/15 °C day/night temperatures and a CO₂,concentration of 340 µmol mol^–1 During the firsthalf of the experimental period the C680 plants grew fasterthan their C340 counterparts so that by week 3 they were twicethe weight this 2 1 superiority in weight persisted until theend of the experiment The faster initial growth of the C680plants was based on an approx 70 % increase in leaf numbersand an approx 30 % increase in their individual area Initially,specific leaf area (cm2 g^–1 leaf) was lower in C680 thanin C340 leaves but became similar in the latter half of theexperiment Shoot organ weights, including petioles and stolons,reflected the C680 plant's better growth in terms of photosyntheticsurface Throughout, C680 plants invested less of their weightin root than C340 plants and this disparity increased with timeAcetylene reduction assays showed that nitrogenase activityper unit nodule weight was the same in both C680 and C340 plantsBoth groups of plants invested about the same fraction of totalweight in nodules Nitrogen contents of plant tissues were similarirrespective of growth regime, but C680 expanded leaves containedslightly less nitrogen and their stolons slightly more nitrogenthan their C340 counterparts However, C680 leaves containedmore non-structural carbohydrate Young, unshaded C680 leavespossessed larger palisade cells, packed more tightly withinthe leaf, than equivalent C340 leaves The reason for the C680regime's loss of superiority in relative growth rate duringthe second half of the experiment was not clear, but more accumulationof non-structural carbohydrate, constriction of root growthand increased self-shading appear to be the most likely causes Trifolium repens, white clover, elevated CO₂, elevated temperature, growth, N₂ fixation, leaf structure     

The Growth and Photosynthesis of Seedling Plants of White Clover at Low Temperature

The growth of white clover (Trifolium repens L.) in conditionstypical of April in Southern England (8 °C day/4 °Cnight, 12 h photoperiod of 90 J m^–2 s^–1 visibleradiation) was extremely slow, whether the plants were dependentfor nitrogen on fixation by their root nodules or were suppliedwith abundant nitrate; although growth was slower in the nodulatedplants. The reasons for slow growth were a large root: shootratio and a small leaf area, particularly in the nodulated plants,and a low photosynthetic rate in all plants. The probable effectsof these characteristics on the growth of white clover withgrasses in mixed pastures are discussed. Trifolium repens L, white clover, low temperature, leaf area, photosynthetic rate, nitrogen supply, growth     

The Effect of Defoliation on Carbohydrate, Protein and Leghaemoglobin Content of White Clover Nodules

Single white clover plants grown in pots of Perlite in a controlledenvironment and completely dependent on N₂ fixation were defoliatedto various degrees (46–85 per cent of shoot weight removed).The soluble protein content of nodules declined by about 20per cent and leghaemoglobin content by 50 per cent in the first4–7 d after defoliation but increased again to controllevels as new leaf tissue appeared. In the short term (2–3h) carbohydrate content of nodules declined to different extentsdepending on the severity of defoliation. The initial declinein N₂ fixation and the respiration associated with it, appearednot to be related to the instantaneous carbohydrate contentof nodules but rather to the supply of current photosynthatefrom the shoot. After 24–48 h, however, the carbohydratecontent of nodules had declined to low levels, regardless ofthe severity (46 or 71 per cent shoot removed) of defoliation.As new leaf tissue appeared carbohydrate levels in all partsof the plant gradually recovered towards control levels. Microscopic examination of nodule sections indicated that onlyafter very severe defoliation (80–85 per cent shoot removed)was nodule deterioration evident. Even here, as the plant establishednew leaves, the damage to nodules was repaired and no noduleloss was apparent. Trifolium repens, white clover, defoliation, carbohydrate, protein, leghaemoglobin     

The Production of Leaves and Stolon Branches on Established White Clover Cuttings in Relation to Temperature and Soil Moisture in the Field

Successive sets of cuttings of three white clover genotypeswere raised in a 15 °C growth room and transferred to thefield at 14 d intervals over the course of a year. Rates ofleaf appearance (leaves per stolon growing point per unit time)were found to be closely correlated with 10 cm soil temperatures.Petiole lengths, and weights of the lamina+petiole increasedin May and decreased towards the end of August, but also exhibiteda marked response to a mid-season water deficit. In the conditionsof the experiment (i.e. in the absence of competition from neighbouringplants) the vast majority of axillary buds developed into visiblebranches at all times of the year. There was, however, an increasein the nodal age at which bud development was first observedin winter. Deferred bud development was also observed, particularlyin one genotype, during periods characterized by dry soil surfaceconditions. The results are discussed in relation to observedpatterns of stolon branching in sward conditions. White clover, Trifolium repens, axillary bud development, branching, growing points, leaf appearance rate, petiole length, soil moisture, soil temperature     

The Growth and Carbon Allocation Patterns of White Clover (Trifolium repens L.) Plants of Contrasting Branching Structure

The growth, morphology and carbon allocation patterns of F1progeny white clover (Trifolium repens L.) plants selected foreither low (‘LBF’) or high (‘HBF’) frequencyof stolon branching were compared in two controlled-environmentexperiments. Selections from within both a small-leaved (‘GrasslandsTahora’) and a large-leaved (‘Grasslands Kopu’)clover cultivar were compared, and plants were grown under arelatively lenient defoliation treatment (expt 1) or under threelevels of defoliation seventy (expt 2). Carbon allocation patternswere measured by ¹⁴CO₂ pulse-chase labelling using fully unfoldedleaves on the main (parent) stolon. LBF and HBF displayed consistent differences in the selectedcharacter though, within cultivars, the difference between selectionswas most pronounced for Kopu. The selections developed fundamentallydifferent branching structures resulting from differences inbranching frequency, with total branch weight per plant averaging122 mg for LBF and 399 mg for HBF (mean of both experiments).More C moved from parent stolon leaves to branches in HBF thanin LBF (mean 22.6% vs. 15.1% respectively of the ¹⁴C exportedfrom source leaves). More C also moved to stolon tissue in HBF,but, counterbalancing this and the difference in allocationto branches, less moved to developing leaves and roots on theparent stolon itself compared to LBF. However, the total weightof developing leaves and roots per parent stolon was generallygreater in HBF than in LBF, probably reflecting greater C importby these sinks from the higher number of branches present perplant in the former selection. HBF plants were consistentlylarger at harvest than LBF plants. There were no defoliationtreatment x selection interactions in C allocation patternsin expt 2. The implications of the results for plant performancein grazed pastures are discussed. Branching, carbon translocation, defoliation, growth, morphology, Trifolium repens, white clover     

The Effect of Lenient Defoliation on the Nitrogen Economy of White Clover: The Contribution of Mineral Nitrogen to Plant Nitrogen Accumulation During Regrowth

Single plants of white clover (Trifolium repens L.) were grownfrom stolon cuttings rooted in sand. All plants were inoculatedwith Rhizobium trifolii, and for 14 weeks received nutrientsolution containing 0.5 mg N each week, as either ammonium ornitrate. Plants were then leniently defoliated or were leftintact and a ¹⁵N-labelled N source was applied at intervalsof 4 d to replace the unlabelled N. Lement defoliation removedfully expanded leaves only; the remaining immature leaves accountedfor 39–44% of the total. At harvests over the following21 d, leaf numbers were counted and dry matter (DM), N contentsand ¹⁵N enrichments of individual plant organs were determined. Rates of leaf emergence and expansion were accelerated in defoliatedplants; numbers of young leaves were similar in defoliated andintact plants. Total DM and N content were less in defoliatedthan intact plants and were not affected by form of N supplied.DM of young leaves, growing points and stolons and N contentof young leaves were, however, greater when ammonium ratherthan nitrate N was supplied. Rates of increase in the contentof plant total N were 8.2 ± 1.36 mg N d^-1 and 10.2±1.82 mg N d^-1 in defoliated and intact plants respectively.The increases were predominantly due to N₂ fixation, since recoveryof ¹⁵N showed that less than 1% of the increment in plant totalN was assimilated mineral N. Nevertheless, the contributionof mineral N to plant total N was 50% more in defoliated thanin intact plants; higher amounts of mineral N were found particularlyin young leaves and growing points. Partitioning of mineralN to nodulated roots increased over time and was greater whenammonium rather than nitrate N was present. White clover, Trifolium repens L. cv. S184, lenient defoliation, N accumulation, N₂ fixation     

Direct Somatic Embryoid Formation on Immature Embryos of Trifolium repens, T. pratense and Medicago sativa, and Rapid Clonal Propagation of T. repens

By direct somatic embryogenesis in vitro a clone of asepticplantlets can be raised from a single immature embryo of Trifoliumrepens (white clover) within about 6 weeks of pollination. Embryoidsare induced directly from intact zygotic embryonic tissue ona culture medium containing 0·025 or 0·05 mg 1^–1BAP and 1·0 g 1^–1 yeast extract. Similar directsomatic embryogenesis has also been achieved for Trifolium pratense(red clover) and Medicago sativa (lucerne). Applications ofembryo propagation by direct somatic embryogenesis are discussed,particularly in relation to multiple screening of host genotypesfor analysis of host/pathogen and legume/Rhizobium interactions. Trifolium repens L., Trifolium pratense L., Medicago sativa L., clover, lucerne, tissue culture, embryoid, somatic embryogenesis, legumes     

The Carbon Economy of Clonal Plants of Trifolium repens L.

Fluxes of carbon between sources and sinks were quantified forclonal plants of Trifolium repens L. (cv. Blanca) in two glasshouseexperiments. Carbon sources were (a) leaves on the parent (=main)stolon apex, or (b) leaves on either young or old branches,and the major sinks of interest were the parent stolon apex,branches, and the adventitious root arising at the same parentstolon node as a young source branch. Defoliation treatmentswere applied to the parent stolon and/or branches (excludingsource branches). Carbon moved freely from the parent stolon to branches and vice-versa;these bidirectional exchanges of C provided important supplementarysources of carbohydrate for the sinks and buffered them againstthe effects of defoliation. Young branches exported more C tothe parent plant (mean=6.3µmol d^–1) than they importedfrom leaves on the parent stolon (5·2µmol d^–1)which, in turn, exceeded the amount fixed by leaves on the branchand utilized within the branch itself (2·7µmold^–1). In contrast, the C economy of old branches was largelyself-contained with, on average, 25·4µmol d^–1exported to the parent plant, 1·8µmol d^–1imported from the parent, and 63·0µmol d^–1fixed and utilized by the branch itself. Thus the growth ofyoung branches was immediately reduced by removal of parentstolon leaves, but old branches were unaffected. An estimated 42% of the C utilized by the main stolon apex originatedfrom branches, while by far the largest proportion (84%) ofthe C used for growth of young nodal roots originated from theassociated branch and not from leaves on the parent stolon towhich the root was directly attached. Key words: Trifolium repens, clonal growth, carbon economy, physiological integration, defoliation     

The Effect of Water Stress on Floral Characters, Pollination and Seed Set in White Clover (Trifolium repens L.)

White clover plants were subjected to either short-term developingwater stress or long-term stable levels of water deficit. Thehort-term stress reduced plant water status to about –2·0MPa over 16 d. The long-term stress was less severe, but wassustained for several weeks. Long-term water stress promotedthe production of inflorescences. However, water stress alsoincreased floret abortion and the premature death of whole flowerheads. The number of ovules per floret was decreased by waterstress. The most striking effect of both long- and short-term waterdeficit was to reduce pollen viability measured with the fluorochromaticassay. This was not an artefact of assay conditions. The pollenfrom water-stressed flower heads was not reversibly dehydrated;it did not score at similar viability to controls after incubationin conditions which hydrate pollen. In addition, the pollenfrom water-stressed plants lost viability more rapidly thanpollen from well-watered plants after removal from the flowerhead. The consequences of reduced pollen viability on seed set wereinvestigated by hand-crossing within and between groups of plantsmaintained for several weeks at three levels of water supply.Flower heads pollinated with pollen from water-stressed plantsset fewer seeds per floret than those pollinated with controlpollen. Key words: Trifolium repens, white clover, water stress, floral characters, seed set     

Effect of Previous Defoliation Regime and Mineral Nitrogen on Regrowth in White Clover Swards: Photosynthesis, Respiration, Nitrogenase Activity and Growth

Small swards of white clover (Trifolium repens L.) cv. Haifawere grown in solution culture in a controlled environment at24 °C day/18 °C night and receiving 500 µE m^-2S^–1 PAR during a 14-h photoperiod. The swards were cuteither frequently (10-d regrowth periods) or infrequently (40-dregrowth) over 40 d before being cut to 2 cm in height. Halfof the swards received high levels of nitrate (2–6 mMN in solution every 2 d) after defoliation while the othersreceived none. Changes in d. wt, leaf area and growing pointnumbers were recorded over the following 10 d. CO₂ exchangewas measured independently on shoots and roots and nitrogenase-linkedrespiration was estimated by measuring nodulated root respirationat 21% and 3% oxygen in the root atmosphere. There was a general pattern in all treatments consisting ofan initial d. wt loss from roots and stubble and reallocationto new leaves, followed by a period of total d. wt gain andrecovery, to a greater or lesser extent, of weight in non-photosyntheticparts. Frequently cut swards had a smaller proportion of theirshoot d. wt. removed by cutting and had a greater shoot d. wt,growing point number and leaf area at the start of the regrowthperiod. As a result of these differences, and also because ofdifferences in relative growth rates, frequently cut swardsmade more regrowth than infrequently cut. Initial photosyntheticrates were higher in frequently cut swards, although the laminaarea index was very low, and it was concluded that stolons andcut petioles made a significant contribution to carbon uptakeduring the first few d. Infrequently cut swards continued toallocate carbon to new and thinner leaves at the expense ofroots and stubble for longer than frequently cut swards andas a result achieved a similar lamina area index after 10 d. Nitrogenase-linked respiration was low in all treatments immediatelyafter cutting: frequently cut swards receiving no nitrate maintainedhigh nitrogenase activity, whereas recovery took at least 5d in infrequently cut swards. Swards which received nitrateafter cutting maintained only low rates of nitrogenase-linkedrespiration and their total nodulated root respiration overthe period was lower than those receiving no nitrogen: greaterregrowth in nitrate fed swards over the 10 d compared to N₂-fixingswards was in proportion to this lower respiratory burden. White clover (Trifolium repens L.), defoliation, regrowth, nitrogen, photosynthesis, respiration, nitrogenase-activity     

Effects of Drought and Defoliation on Bud Viability in Two Caespitose Grasses

Axillary bud number, bud respiratory activity, and photosyntheticcanopy re-establishment after defoliation were determined fortwo bunchgrass species, Agropyron desertorum and Agropyron spicatum,which were exposed to draughted, natural or irrigated conditions.These field treatments were repeated annually on the same plantsfor the period 1984–1986. Bud respiratory activity wasexamined using the tetrazolium test, which was validated withthe vital stain Evan's blue, at the end of the study In spring of the third year, the number of axillary buds ontillers of both species was lowest in the drought treatment.Most of these buds, and those observed immediately after defoliationin 1985, were metabolically active. These results indicate thatafter mid-season defoliation under drought, when no re-growthoccurred, the re-growth capacity was not limited by bud numberor viability. After 3 years of defoliation, tiller number andgrowth in both species were reduced in the following springunder all water regimes. This reduction was present 1 year earlierin the drought treatment than in the treatments with higherwater availability. Permanent dormancy or death of the replacementaxillary meristems can explain this plant response. Continueddefoliation of the tillers under drought would reduce the photosyntheticarea further, and probably affect the persistence of these speciesin the community Agropyron desertorum, Agropyron spicatum, crested wheatgrass, bluebunch wheatgrass, drought, defoliation, re-growth, bud viability, tetrazolium, Evan's blue     

Leaf and Tiller Production of Prairie Grass (Bromus willdenowii Kunth) and Two Ryegrass (Lolium) Species

A detailed morphological study of three prairie grass cultivars(Bromus willdenowii Kunth) was conducted under ‘vegetative’and ‘reproductive’ growth conditions (short andlong photoperiods) and at different temperatures. Perennialryegrass (Lolium perenne L.) and Westerwolds ryegrass (Loliummuhiflorum Lam.) were compared during vegetative growth. Prairie grass had higher leaf appearance rates (leaves per tillerper day) and lower site filling (tillers per tiller per leafappearance interval) than the ryegrass species. Tillering rates(tillers per tiller per day) were also lower, except under vegetativeconditions at 4C. Low tiller number in prairie grass was notdue to lack of tiller sites but a result of poor filling ofthese sites. Lower site filling occurred because of increaseddelays in appearance of the youngest axillary tiller and lackof axillary tillers emerging from basal tiller buds. In prairiegrass, no tillers came from coleoptile buds while only occasionallydid prophyll buds develop tillers. Low tiller number in prairiegrass was compensated for by greater tiller weight. Prairiegrass had more live leaves per tiller, greater area per leafand a high leaf area per plant. Considerable variation between cultivars was found in prairiegrass. The cultivar ‘Bellegarde’ had high leaf appearance,large leaves and rapid reproductive development, but had lowlevels of site filling, tillering rate, final tiller numberand herbage quality during reproductive growth. ‘Primabel’tended to have the opposite levels for these parameters, while‘Grasslands Matua’ was intermediate and possiblyprovided the best balance of all plant parameters. prairie grass, Bromus willdenowii Kunth, perennial ryegrass, Lolium perenne L., Westerwolds ryegrass, Lolium multiflorum Lam., temperature, photoperiod, leaf appearance, leaf area, tillering, site filling, tillering sites, yield     

The Influence of Irradiance Distribution on the Growth of White Clover (Trifolium repens L.) in Differently Managed Canopies of Permanent Grassland

Growth of white clover was investigated in permanent grasslandcut three or five times per year. The influence of cutting frequencyand nitrogen fertilization on dry-matter yield, leaf-area distributionand the distribution of photosynthetically active radiationwithin the canopy were examined. In the five cut treatments, total dry-matter yield was nearlyequal, with and without nitrogen. However, nitrogen practicallyeliminated white clover. Leaf-area distribution showed characteristicpatterns for the different treatments. The small proportionof white clover in the treatment with nitrogen fertilizationwas thought to be due to the large leaf area of the other speciesat heights which white clover could not attain. This conclusionwas supported additionally by the radiation measurements withinthe canopy. The sunlit fractional area within canopy layers was measuredwith ‘quantum sensors’ and calculated from canopytransmission measured with tube solarimeters. The leaf areaindex of white clover was highly correlated (r² = 0.68) withthe sunlit fractional area above the canopy layers where whiteclover was present. This response of white clover leaf growth to the light regimeis discussed in relation to the potential petiole growth. White clover, Trifolium repens L., permanent grassland, irradiance distribution, sunlit fractional area, petiole extension, leaf area, dry matter, stratified clipping     

Oxygen Supply Limits Nitrogenase Activity of Clover Nodules After Defoliation

The aim of this study was to test the effect of oxygen partialpressure as a possible limiting factor of nitrogen fixationfollowing defoliation. The response of nitrogenase activity(C₂H₂-reduction) of defoliated and undefoliated white and redclover plants (Trifolium repens L. and Trifolium pratense L.)to either 19 kPa oxygen or 55 kPa oxygen was investigated. Priorto defoliation, white clover plants were grown for five weeksin a growth chamber, and red clover plants for 7 or 11 weeksin a glasshouse. The results included measurements of ¹⁶N₂-uptake. Increasing oxygen partial pressure from 19 to 55 kPa severelyrestricted nitrogenase activity of undefoliated white cloverplants; however, 2 h after complete defoliation, the same treatmentcaused a significant increase. A fivefold increase in nitrogenaseactivity upon exposure to the elevated oxygen partial pressurewas found at the end of a 24 h period. This beneficial effectdecreased gradually from 1 to 5 d after defoliation. The responseof recently defoliated red clover plants to 55 kPa oxygen partialpressure was similar to that of white clover, independentlyof plant age. The gradual recovery of nitrogenase activity duringthree weeks of regrowth was associated with a simultaneous changein the response to increased oxygen partial pressure, leadingagain to the response of undefoliated plants. These data suggested that lack of oxygen at the site of nitrogenfixation, resulting from a dramatic increase in oxygen-diffusionresistance, is the main factor limiting nitrogenase activityfollowing defoliation. Trifolium repens L., Trifolium pratense L., white clover, red clover, defoliation, regrowth, nodules, nitrogen fixation, nitrogenase activity, oxygen limitation