Aspects of the Comparative Physiology of Ranunculus bulbosus L. and Ranunculus repens L.: II. Carbon Dioxide Assimilation and Distribution of Photosynthates

Detailed analysis of the interrelationships between sourcesof photosynthate production and sites of utilization in thetaxonomically closely related species Ranunculus bulbosus L.and R. repens L. showed that leaves whether present on rosette,stem, or stolon had similar levels of ¹⁴CO₂-fixation but thepattern of distribution of radiocarbon to the rest of the plantdiffered. Fruits of R. bulbosus had a lower fixation rate thanleaves but were characterized by total retention of the fixedradiocarbon. Rosette leaves of R. bulbosus supplied the youngleaves, developing apices in the rosette, roots, and corms,whereas the labelled assimilates from cauline leaves were evenlydistributed between reproductive and vegetative parts. The cormwas the major sink both at the flowering and fruiting stages.When plants were treated with ¹⁴CO₂ in the field even higherlevels of radiocarbon moved into the corm than in comparableexperiments under greenhouse conditions. The rosette leaf ofR. repens exported mainly to actively growing stolons in plantswith many stolons bearing rooted ramets although growth of astolon was also substantially supported by photosynthates producedby its own ramets. A proportion of the radiocarbon fixed byleaves of mature ramets was exported and moved in a predominantlyacropetal direction into the stolon apex, stolon axis, and youngramets of the same stolon. The stock in R. repens had a muchlower demand for assimilates than the corm in R. bulbosus. The results are consistent with the concept that R. bulbosusoperates a conservative policy involving the replacement ofthe parent in situ by a daughter from the corm, coupled withextensive fruit production. In R. repens the emphasis is onlateral spread and exploitation of substantial areas of groundby vegetative spread and replacement of the parent by daughtersmany of which may occupy sites some distance from the parent.     

Seasonal Allocation and Efficiency Patterns of Biomass and Resources in the Perennial Geophyte Sparaxis grandiflora Subspecies fimbriata (Iridaceae) in Lowland Coastal Fynbos, South Africa

The seasonal allocation and efficiency of biomass and resources(starch, soluble carbohydrates, N, P, K, Ca, Mg, Na, Zn, Fe,Mn and Cu) within and between the constituent plant parts ofSparaxis grandiflora subspecies fimbriata , a deciduous, synanthousgeophyte, are described. The sequential production of roots,leaves and a new daughter corm, with fruit production phasedin during the development of the daughter corm, results in continualtransfer of resources when the plant is not dormant. The newdaughter corm serves as the major sink for the allocation ofdry matter and resources during the vegetative, reproductiveand senescent periods. The parent corm is the major source ofresources in the early stages of the vegetative period. Theamount of resources allocated to the reproductive structureswas comparatively low when compared with that allocated to vegetativeplant parts. The corms have highly flexible storage capacitiesfor a variety of nutrients, particularly the better conservednutrients such as nitrogen, phosphorus, potassium, and carbohydrates.The dynamic patterns for the majority of the resources revealedhigh allocation when metabolic activity was also highest. Efficientrecovery or recycling of important resources from senescingorgans results in a similar or greater allocation to the daughtercorm than the amount that was present in the parent corm ofthe previous growth season. This is seen as an advantage tothese plants that grow in a seasonal environment where soilsare of low nutrient status.Copyright 1994, 1999 Academic Press Dry mass, efficiency, phenophase, resource allocation, Sparaxis grandiflora, fynbos     

Mobilization of nitrogen reserves during regrowth of defoliated Trifolium repens L. and identification of potential vegetative storage proteins

Although it is well established that carbon reserves contributeto shoot regrowth of leguminous forage species, little informationis available on nitrogen reserves except in Medicaqo sativaL. and Trifolium subterraneum L. In this study, reserves werelabelled with ¹⁵N to demonstrate the mobilization of endogenousnitrogen from roots and stolons to regrowing leaves and newstolons during 24 d of regrowth in white clover (Thfolium repensL.). About 55% and 70%, respectively, of the nitrogen contentsof these organs were mobilized to support the regrowth of leaves.During the first 6 d, nitrogen in regrowing leaves came mainlyfrom N reserves of organs remaining after defoliation. Afterthese first 6 d of regrowth, most of the shoot nitrogen wasderived from exogenous nitrogen taken up while the contributionof nitrogen reserves decreased. After defoliation, the buffer-solubleprotein content of roots and stolons decreased by 32% duringthe first 6 d of regrowth. To identify putative vegetative storageproteins, soluble proteins were separated using SDS-PAGE ortwo-dimensional electrophoresis. One protein of 17.3 kDa instolons and two proteins of 15 kDa in roots seemed to behaveas vegetative storage proteins. These three polypeptides, initiallyfound at high concentrations, decreased in relative abundanceto a large extent during early regrowth and then were accumulatedagain in roots and stolons once normal growth was re-established. Key words: White clover, regrowth, ¹⁵N-labelled, vegetative storage proteins, electrophoresis     

Effects of Soil Nutrients and Moisture on Root/Shoot Ratios in Lolium perenne L. and Trifolium repens L

The dry weights of foliage and roots (excluding stembases) werecompared in a 3³x2² factorial experiment with Lolium perenneL. and Trifolium repens L. grown in pots with three nitrogen(0–300 ppm N), phosphorus (0–300 ppm P), and soilmoisture (pF 3.2, pF 2.4, and waterlogged) treatments, eachduplicated. Root/shoot ratios were influenced significantly (P < 0.001)by N, P, and W (soil moisture), but these main effects werequalified by highly significant first-order interactions andsecond-order interactions which included N and P. The lowestR/S ratios occurred where N, P, and W were all non-limiting,and the highest where there was an imbalance of these factors. The explanation suggested is that a decrease in the availabilityof N, P, or W causes an increase in the relative weight of roots,but in the presence of N, P, or W deficiency an increase inany one of these factors causes a further increase in the R/Sratio by increasing the relative efficiency of the foliage.     

Field evidence of plastic growth responses to habitat heterogeneity in the clonal herbRanunculus repens

Stolon internode lengths were measured on plants of the clonal herbRanunculus repens growing in a hay meadow which was subject to disturbance by mole (Talpa europaea) activity. Within the site three habitat types were recognized: closed grassland, the open ground of fresh molehills and the grass-molehill boundary. The lengths of stolon internodes ofR. repens differed significantly in each of the three habitats. The shortest internodes occurred on stolons on the open molchills. The longest occurred in the closed grassland habitat. The type of habitat in which parent ramets were rooted did not significantly influence the length of internodes on their daughter stolons. The length of a stolon internode was determined by its immediate surrounding habitat type. Consecutive internode lengths on a given stolon showed considerable plasticity, shortening significantly as stolons spread onto molehills from surrounding habitats, and increasing significantly as stolons advanced from a molehill into the surrounding closed grassland habitat. These results are consistent with the proposition that under favorable conditions (on the molehills, where resources are expected to be more abundant, and competition absent) internode lengths shorten and the plant forages intensively, whereas under conditions of low resource availability (in the closed grassland, where competition occurs) internode lengths increase, allowingR. repens to forage extensively. Such morphological plasticity may promote more efficient exploitation of resource-rich sites and more rapid vacation of resource-poor sites.     

Vascular Architecture of a Large-leafed Genotype ofTrifolium repens

The objectives of this study were to identify the vascular connectionsfrom roots to upper axial bundles in one genotype ofTrifoliumrepensL. ‘Grasslands Kopu’, identify pathways followedby the transpiration stream, and establish whether these pathwayscould account for previously-observed patterns of clonal integration.The study provides new information on vascular connections betweenroot and parent and branch stolons at nodes possessing botha root and a branch, and to the first two leaves on branch stolons.A nodal root is connected to the lower nearside axial bundleof the parent stolon but to both lower and upper nearside axialbundles of the branch. Upper sympodia provide a long-distancetransport pathway from a parent stolon to the apex of branchstolons. Lower sympodia are functionally different, providingshort-distance transport to structures in close proximity tothe source root. This is consistent with observed patterns ofclonal integration inT. repensand may provide a simple architecturalmechanism facilitating foraging.Copyright 1998 Annals of BotanyCompany Acid fuchsin, clonal integration, foraging, physiological integration, serial sections, white clover,Trifolium repens(L.), vascular architecture, xylem transport.     

Evidence For Root Contraction In White Clover (Trifolium repens L.)

White clover (Trifolium repens L.) stolons become buried inthe field. It was observed that this also occurred in the greenhousewhere the accepted mechanisms of burial, treading by livestockand earthworm casting, did not occur. It was also observed thatthe crown of seedling T. repens plants become closely appressedto the soil. Experiments showed that, regardless of varietyof T. repens or depth of planting, all seedling hypocotyls firstlift the cotyledons clear of the soil, then ‘ contract’towards the soil until the cotyledons are in contact with orbelow the soil surface. Auxanometers were used to measure therate and extent of this contraction and were also attached tostolon nodes in experiments which showed that stolons move downwardsrelative to the soil surface and that the speed and extent ofthis duration varied with soil type. A further experiment showedthat only rooted nodes show this behaviour. The force exertedby the contraction of nodal roots was estimated experimentallyas 0.21 N g^-1fresh root. A mechanism for the root contraction,based on examination of root anatomy of seedling tap-roots andnodal roots, is suggested. These experiments provide evidencefor root contraction in T. repens which may lead to stolon burial.The importance of this to T. repens as a pasture species andas a means of further improving T. repens varieties is discussed.Copyright 1999 Annals of Botany Company White clover, Trifolium repens, L., stolon, seedling, burial, root, nodes, nodal, force, contractile, soil resistance, pasture, phloem, fibres.     

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     

Phosphate Uptake and Transport by Roots and Stolons of Intact White Clover (Trifolium repens L.)

The extent to which phosphate can be absorbed directly fromthe outer medium by stolon internodes and contribute to thetotal accumulation of phosphate by intact plants of white clover(Trifolium repens L. cv. Blanca) was assessed in hydroponicexperiments in a controlled environment room. The uptake ofphosphate by intact roots or stolons was measured by sealinga segment (6-0 mm long) across a flow-cell in which ³²P-labellednutrient solution was circulated for 24 h, the rest of the rootsystem receiving unlabelled nutrient solution. The rate of uptakeof phosphate (µmol g^–1 d^–1 dry wt. basis)by roots was more than 300 times that by intact stolons. Pretreatmentof stolons by gentle abrasion to remove cuticle, so as to simulatethe condition of stolons in the field, increased the uptakeof phosphate 7-fold compared with that of intact stolons. However,the potential of stolons to contribute to the P status of whitedover in the field was calculated to be small (5%). When an incision was made through the hypodermal layer of stolons,the rate of phosphate uptake greatly increased, attaining 71%of that by root segments. This increase, which was greater athigher phosphate concentrations, indicates that the suberi.zedhypodermis constitutes a major barrier to the influx of phosphatein the stolon. After withholding phosphate for different time intervals, thesubsequent rate of phosphate uptake by roots was increased 2-3-foldafter 2 d phosphate deprivation and 3-4-fold after 6 d or 13d phosphate deprivation. A higher proportion of absorbed phosphatewas transported to shoots in phosphate-deprived plants. After1 d of uptake following restoration of the phosphate supply,the concentrations of labelled phosphate in shoots were greaterthan in control plants, although the concentrations of labelin roots was less. However, the rate of uptake of phosphateby stolons, following deprivation, was not significantly increased.These results suggest that the mechanism regulating the enhancedrate of phosphate loading into the xylem, initiated by a periodof phosphate deprivation, is specific to roots and is not inducedin stolons. The results are discussed in relation to the growth and acquisitionof phosphate by white clover in the field. Key words: Nutrient deficiency, phosphate, stolons, transport (ions), Trifolium repens     

Evaluating contemporary pollen dispersal in two common grassland species Ranunculus bulbosus L. (Ranunculaceae) and Trifolium montanum L. (Fabaceae) using an experimental approach

Pollen flow is a key biological process that connects plant populations, preventing genetic impoverishment and inbreeding. Pollen‐mediated long‐distance dispersal (LDD) events are especially important for plant species in increasingly fragmented landscapes. Patterns of pollen dispersal were directly estimated and dispersal kernels modelled in an experimental population of Ranunculus bulbosus and Trifolium montanum to determine the potential for LDD. Eight and 11 microsatellite markers were used for R. bulbosus and T. montanum, respectively, to run a likelihood‐based paternity analysis on randomly chosen offspring (N_total = 180 per species) from five maternal plants. High rates of selfing were found in R. bulbosus (average 45.7%), while no selfing was observed in T. montanum. The majority (60%) of mating events occurred at very short distances: the median of the observed dispersal distances was 0.8 m in both species, and the average distances were 15.9 and 10.3 m in R. bulbosus and T. montanum, respectively. Modelling the pollen dispersal kernel with four different distribution functions (exponential‐power, geometric, 2Dt and Weibull) indicated that the best fit for both species was given by a Weibull function. Yet, the tail of the T. montanum pollen dispersal kernel was thinner than in R. bulbosus, suggesting that the probability for LDD is higher in the latter species. Even though the majority of pollen dispersal occurred across short distances, the detection of several mating events up to 362 m (R. bulbosus) and 324 m (T. montanum) suggests that pollen flow may be sufficient to ensure population connectivity in these herb species across fragmented grasslands in Swiss agricultural landscapes.     

Accumulation, Partitioning and Redistribution of Dry Matter and Mineral Nutrients in Ixia flexuosa L., with Special Reference to its Cormaceous Habit

The seasonal dynamics of uptake, partitioning and redistributionof dry matter, N, P, K, S, Ca, Mg, Na, Cl, Fe, Zn, Mn and Cuby the cormaceous plant Ixia flexuosa were studied in pot cultureat Perth, Western Australia. Dry matter and P, N, K, Zn andCu were redistributed from the mother corm with about 90 percent net efficiency: there was no net redistribution of Ca,Na, Fe or Mn. The efficiency of redistribution from the leafyshoot to fruits and the new season's corm was 80 per cent forN and P, 24–49 per cent for K, Cu and Zn, and 0–15per cent for Na, Fe, Ca, Mn, Cl, Mg, S and dry matter. Redistributionfrom the mother corm and vegetative organs could have suppliedthe replacement corm, cormlets and fruits with 32–53 percent of their S, K, P, N, Cu and Zn, and 11–25 per centof their Ca, Cl, Mn, Mg and dry matter. The mature replacementcorm had over 60 per cent of the plant's N and P, 25–50per cent of its dry matter, Zn, Cu, Mg, K and Cl, but less than20 per cent of its Ca, Na, Fe and Mn. Each plant produced anaverage of 12 cormlets; these had 35 per cent of the dry matterand 23–47 per cent of the amount of a particular nutrientin the new season's corms. Fruits had less than 16 per centof the dry matter and each mineral in the mature plant. Ratesof mineral intake by Ixia were much lower than reported forcrop plants, and may be related to the long growing season ofthe species. Ixia polystachya L., corm, nutrition, mineral nutrients, nutrient redistribution     

Seasonal Dynamics of the Accumulation, Distribution and Redistribution of Dry Matter and Mineral Nutrients in a Weedy Species of Gladiolus (Gladiolus caryophyllaceus)

The seasonal dynamics of the accumulation, distribution andredistribution of dry matter and 12 mineral nutrients by a weedyspecies of gladiolus (Gladiolus caryophyllaceus) were studiedat Perth, Western Australia, where it has colonized the nutrient-poorsandy soils. Parent corms sprouted in autumn, and the plantshad completed their growth cycle by early summer. The maturereplacement corm had 15-25% of the plant's P, Ca, Na, Zn andCu, 5-15% of its K, N, Cl, Mg, S and dry matter, and < 5%of its Fe and Mn. Seeds had 26% of the plant's dry matter, 60%of its N and P, 21-33% of its S, Mg, Cu and K, 5-20% of itsFe, Mn and Zn, and < 5% of its Ca and Na. The mature vegetativeshoot had 47% of the plant's dry matter and over 40% of eachnutrient, except for N, P and Cu. Phosphorus, K and N were redistributedfrom the parent corm with over 85% efficiency, S, Mg, Zn andCu with 60-70% efficiency, but there was < 10% redistributionof Ca, Na, Cl, Fe and Mn. The efficiency of redistribution fromthe leafy shoot was over 70% for N and P, 29-52% for K, Mg andCu, 16-20% for S, Zn and Cl, but negligible for Ca, Na, Fe andMn. Redistribution from the shoot could have provided the replacementcorm and seeds with 53-98% of their Cu, Mg, N, P and K, and29-38% of their S, Zn and dry matter. Seeds contained over 60%of each nutrient in a capsule, except for Ca, Na and Fe. Redistributionfrom the capsule walls could have provided 13-19% of the P,Cu and Zn, and 3-7% of the N, K, Mg and dry matter accumulatedby seeds. Each plant produced an average of 520 seeds. Removalof flowers and buds at first anthesis resulted in a larger replacementcorm containing a greater quantity of most nutrients, indicatingcompetition between the replacement corm and seeds for nutrients.Redistribution from parent to replacement cormlets in the absenceof shoot and root development was high, with over 50% of thedry matter and each nutrient, except for Ca, being transferred.Concentration of nutrients were low in all organs of G. caryophyllaceus,especially the replacement corm. It was concluded that the effectiveredistribution of key nutrients, such as N and P, to reproductivestructures and tolerance of low internal concentrations of nutrientscontribute to the capacity of G. caryophyllaceus to colonizeand persist on infertile soils.Copyright 1993, 1999 AcademicPress Gladiolus caryophyllaceus, corm, distribution, dry matter, gladiolus, mineral nutrients, nutrient accumulation, nutrient redistribution, seasonal growth, weed     

Assimilate Partitioning in Red and White Clover Either Dependent on N2 Fixation in Root Nodules or Utilizing Nitrate Nitrogen

During vegetative growth in controlled environments, the patternof distribution of ¹⁴C-labelled assimilates to shoot and root,and to the meristems of the shoot, was measured in red and whiteclover plants either wholly dependent on N₂ fixation in rootnodules or receiving abundant nitrate nitrogen but lacking nodules. In experiments where single leaves on the primary shoot wereexposed to ¹⁴CO₂, nodulated plants of both clovers generallyexported more of their labelled assimilates to root (+nodules),than equivalent plants utilizing nitrate nitrogen, and thiswas offset by reduced export to branches (red clover) or stolons(white clover). The intensity of these effects varied with experiment.The export of labelled assimilate to growing leaves at the terminalmeristem of the donor shoot was not influenced by source ofnitrogen. Internode elongation in the donor shoot utilized nolabelled assimilate. Whole plants of white clover exposed to ¹⁴CO₂ on seven occasionsover 32 days exhibited the same effect on export to root (+nodules),which increased slightly in intensity with increasing plantage. Nodulated plants had larger root: shoot ratios than theirequivalents utilizing nitrate nitrogen. Trifolium repens, Trifolium pratense, red clover, white clover, nitrogen fixation, nitrate utilization, assimilate partitioning     

The Use of a Closed System Flow-Through Enclosure Apparatus for Studying the Effects of Partial Pressure of Dinitrogen in the Atmosphere on Growth of Trifolium repens L. and Lolium perenne L.

A closed system flow-through enclosure apparatus was constructedand used to enclose mixtures ofLolium perenne L. (perennialryegrass cv. Trani) and nodulatedTrifolium repens L. (whiteclover cv. Blanca) growing in soil in pots. There were no significantdifferences between the shoot growth, in terms of dry matteraccumulation and nitrogen content, of mixtures in the systemcompared to that of mixtures grown in a standard growth cabinet.This demonstrated that stable ambient conditions could be achievedby the closed system and its control circuits and that therewere no apparent side-effects of the recirculatory gases.Reducingthe partial pressure of dinitrogen in the atmosphere affectedwhite clover, but not perennial ryegrass. A fairly rapid effectwas observed 4 h after reduction in dinitrogen partial pressureas some of the clover leaves folded downwards along the petiole.These same effects were observed at two different partial pressuresof dinitrogen (22% and 39%) and with two different replacementgases (argon and helium). In the longer term (11 to 16 d) drymatter accumulation and nitrogen content of the clover shootswere significantly reduced. These effects of reduced partialpressure of dinitrogen were observed in both nodulated (NOD⁺)and nitrate-dependent (NOD^–) clover. Possible reasonsfor these effects were discussed with particular reference toimpurities in the gases used, stomatal responses and plant waterrelations. It was concluded that the closed system flow-throughapparatus provides a useful tool for studying whole plant-soilsystems and, in particular, the cycling of nitrogen. However,the use of a replacement gas to reduce the cost of labelleddinitrogen was obviously not a viable proposition. Key words: Dinitrogen, flow-through system, Lolium perenne, partial pressure, Trifolium repens     

Effect of Nitrogen Supply on the Grass and Clover Components of Simulated Mixed Swards Grown under Favourable Environmental Conditions I. Carbon Assimilation and Utilization

Simulated mixed swards of Perennial Ryegrass (Lolium perenneL.) cv. S23 and White clover (Trifolium repens L.) cv. S100were grown from seed under a constant 20 °C day/15 °Cnight temperature regime and their growth and carbon economyexamined. The swards received a nutrient solution daily, whichcontained either High (220 mg l^–1) or Low (10 mg l^–1)nitrate N. Rates of canopy photosynthesis and respiration, and final drymatter yields were similar in the two treatments although theproportions of grass and clover differed greatly. The Low-Nswards were made up largely of clover. The grass plants in theseswards had high root: shoot ratios and low relative photosyntheticrates – both signs of N deficiency – and were clearlyunable to compete with the vigorously growing Low-N clover plants.These had higher relative growth rates and dry matter yieldsthan their High-N counterparts. In the High-N swards clovercontributed around 50 per cent to the sward dry weight throughoutthe measurement period despite having a smaller proportion ofits dry weight in photosynthetic tissue (laminae) than grassover much of it. The latter was compensated for, initially bya higher specific leaf area than grass, and later by a higherphotosynthetic rate per unit leaf weight. The results are discussedin relation to observed declines in the clover content of swardsafter the addition of nitrogen fertilizer in the field. Trifolium repens, white clover, Lolium perenne, perennial ryegrass, nitrogen, photosynthesis, carbon balance     

Effects of CO2 enrichment and intraspecific competition on biomass partitioning, nitrogen content and microbial biomass carbon in soil of perennial ryegrass and white clover

Seedlings of perennial ryegrass (Lolium perenne L. cv. Parcour)and white clover (Trifolium repens L. cv. Karina) grown at fivedifferent plant densities were exposed to ambient (390 ppm)and elevated (690 ppm) CO₂ concentrations. After 43 d the effectsof CO₂ enrichment and plant density on growth of shoot and root,nitrogen concentration of tissue, and microbial biomass carbon(C_mic) in soil were determined. CO₂ enrichment of Lolium perenneincreased shoot growth on average by 17% independent of plantdensity, while effects on root biomass ranged between -4% and+ 107% due to an interaction with plant density. Since tilernumber per plant was unaffected by elevated CO₂, the small responseof shoot growth to CO₂ enrichment was atributed to low sinkstrength. A significant correlation between nitrogen concentrationof total plant biomass and root fraction of total plant drymatter, which was not changed by CO₂ enrichment, indicates thatnitrogen status of the plant controls biomass partitioning andthe effect of CO₂ enrichment on root growth. Effects of elevatedCO₂ and plant density on shoot and root growth of Trifoliumrepens were not significantly interacting and mean CO₂-relatedincrease amounted to 29% and 66%, respectively. However, growthenhancement due to elevated CO₂ was strongest when leaf areaindex was lowest. Total amounts of nitrogen in shoots and rootswere bigger at 690 ppm than at 390 ppm CO₂. There was a significantincrease in C_mic in experiments with both species whereas plantdensity had no substantial effect. Key words: CO₂ enrichment, intraspecific competition, biomass partitioning, Lolium perenne, Trifolium repens, grassland     

Seed survival and seasonal emergence in some species of Geranium, Ranunculus and Rumex

Seeds of 14 species were collected, usually in each of 2 or 3 yr, and mixed with the top 7.5 cm of sterilised soil confined in cylinders sunk in the ground outdoors and cultivated three times yearly. Seedling emergence was recorded for at least 3 and usually 5 yr. Most seedlings of Ranunculus bulbosus and Rumex acetosa appeared during the autumn of sowing, while the main emergence of Geranium pratense. Ranunculus acris, Rumex conglomeratus and R. maritimus was in the following spring. Emergence of G. pyrenaicum and G. robertianum took place throughout much of the year after sowing. Seeds of all these species were relatively short-lived in cultivated soil, and few seedlings appeared after the second year. Seed survival was greater in G. dissectum. G. molle, Rumex acetosella. Ranunculus sceleratus and especially R. repens, of which emergence in the fifth year after sowing amounted to 4% of the seeds sown. Emergence from the persistent seed bank was mainly from May to September for G. dissectum and G. molle and during autumn for Ranunculus flammula and R. sceleratus, while seedlings of Ranunculus repens and Rumex acetosella appeared sporadically over most of the year.     

Nutrient Solution pH Control using Dipolar Buffers in Studies of Trifolium repens L. Nitrogen Nutrition

In studies of Trifolium repens nitrogen nutrition, the controlof nutrient solution pH using dipolar buffers, was evaluatedin tube culture under sterile conditions. Five buffers; MES,ADA, ACES, BES and MOPS with pK₂s (20 °C) of 6.15, 6.60,6.90, 7.15 and 7.20 respectively, at a concentration of 2.0mol m^–3, were provided to inoculated Trifolium repensgrowing in nutrient solution containing 7.13 mol m^–3 nitrogenas (NH₄)₂SO₄. Initial pH of each solution was adjusted to theappropriate buffer pK₂ Two buffers, ADA and ACES completelyinhibited plant growth. The remaining buffers had little effectin limiting pH change, although plant dry matter was higherand nodule numbers lower in the presence of these buffers. MESand MOPS were supplied to nutrient solutions with and without7.13 mol m^–3 (NH₄)₂SO₄, at concentrations ranging from0–12 mol m^–3. MES at 9 mol m^–3 and 12 molm^–3 reduced growth of plants reliant on the symbiosisfor providing nitrogen. The provision of MES to plants providedwith NH₄⁺ significantly increased plant yield and reduced nodulenumber at all concentrations. MOPS did not affect plant yieldor nodule number. The use of dipolar buffers in legume nitrogennutrition studies is considered in terms of buffering capacity,and the side effects on plant growth and symbiotic development. Key words: Ammonium, Dipolar buffer, Nitrogen nutrition, pH control, Symbiosis, Trifolium repens     

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     

Patterns of 14C-labelled Assimilate Partitioning in Red and White Clover During Vegetative Growth

A quantitative analysis of the ¹⁴C-labelled assimilate suppliedby the expanded leaves on the primary shoot to growing leaves,stem, lateral shoots (branches or stolons) and roots in redand white clover was conducted during vegetative growth. Stem growth of the primary shoot was inhibited in both cloversand utilized no energy resources. The growing leaves at theprimary shoot apex of white clover imported 4 per cent of theshoot's assimilate compared with 10 per cent in red clover.At the basal end of the primary shoot, the tap root of whiteclover imported 16 per cent of the shoot's assimilate comparedwith 22 per cent in red clover. Branches in red clover and stolonsin white clover were by far the largest sinks for primary shootassimilate, importing 39 per cent and 63 per cent of the labelledassimilate, respectively. Analyses of the translocation of assimilate from individualprimary shoot leaves demonstrated that in both clovers olderleaves exported more of their assimilate to branches or stolons,whereas younger leaves exported more of their assimilate toroots, and possibly in white clover, to growing leaves at thetip of the shoot. Of the labelled assimilate exported to branchesor stolons, each primary shoot leaf exported preferentiallyto the branch or stolon in its own axil, but in addition exportedsubstantial quantities of assimilate to all other axillary shoots,particularly those arising from basal axils where the subtendingleaf had died. Trifolium repens, Trifolium pratense, red clover, white clover, assimilate partitioning, perennation