On the Ordered Development of Plants 2. Self-thinning in Plant Communities

It is shown that the hypothesis that a growing point on a vegetativeplant requires a minimum rate of supply of assimilate to continuegrowth can quantitatively describe self-thinning in communitiesof Trifolium subterraneum. The hypothesis can also be used toexplain the different relationships observed between mean plantweight and plant density when plants are grown in full daylight,70 per cent shade, and transferred between the two light environments.The implications of the hypothesis to self-thinning in naturalplant communities are discussed. Self-thinning, assimilate, plant development     

Relationship between Colour, Phenolic Content and Impermeability in the Seed Coat of various Trifolium subterraneum L. genotypes

The relationship between seed colour, phenol content of thetesta and water impermeability in dark and light seeded genotypesof Trifolium subterraneum L. was investigated. The developmentof water impermeability and catechol oxidase activity in expandingseeds of two genotypes was monitored. The results show catecholoxidase activity decreases as seed colour changes from greento purple, but the potential to become impermeable with dehydrationis not acquired until later in seed development. Both waterpermeable and impermeable seeds of dark coloured genotypes containsubstantial amounts of phenol in the lumen of the Malpighiancells; light coloured testas do not. It is concluded that the darkening of the testa of Trifoliumsubterraneum is associated with oxidation of phenol by catecholoxidase but that development of impermeability is independentof this process. Trifolium subterraneum L., subterranean clover, seed coat impermeability, catechol oxidase, phenol, testa colour     

The Effects of Rootzone Salinity and Hypoxia on Shoot and Root Growth in Trifolium Species

The effect of concurrent salinity (0-60 mM NaCl) and rootzonehypoxia (flooding for up to 15 d) on shoot and root growth andshoot ion concentrations of six species of Trifolium (T. subterraneumL., T. fragiferum L., T. michelianum Savi., T. isthmocarpumBot., T. purpureum Lois., and T. repens L.), was studied intwo greenhouse experiments. There was a significant salinityx flooding effect for shoot yield but no significant salinityx flooding x species interaction although individual speciesdiffered significantly (P < 0·001) in their growthresponse to the saline or flooded conditions separately. Concentrationsof Na and Cl in the shoots of all species increased with increasingperiods of saline flooding and there was a significant salinityx flooding interaction. Sodium and Cl concentrations were significantlyhigher (P < 0·001) in T. purpureum, the species inwhich shoot growth was most depressed by saline flooding, thanother species. In T. michelianum, T. fragiferum and T. repens,fresh and dry weight of roots increased with flooding underboth saline and non-saline conditions while in T. subterraneumroot growth decreased. A significant proportion of the increasedroot growth in the first three species occurred as new adventitiousroots. These roots had higher percentages of internal gas spaceswithin the root tissue even in the presence of salinity comparedwith roots from non-flooded conditions. There were also significantlymore gas spaces in the total root tissue in T. fragiferum andT. repens under saline-flooding than in roots of T. subterraneum.Electron micrographs of the root cross sections illustratedthe presence of these gas spaces or aerenchyma. Trifolium fragiferum, T. repens and T. michelianum are morelikely to be suited to growth in soils prone to high salinityand to flooding than are T. subterraneum, T. purpureum and T.ishmocarpum.Copyright 1993, 1999 Academic Press Trifolium subterraneum, Trifolium fragiferum, Trifolium michelianum, Trifolium isthmocarpum, Trifolium purpureum, Trifolium repens, salinity, flooding, hypoxia, adventitious roots, aerenchyma, subterranean clover, white clover, strawberry clover, purple clover, balansa clover     

Response of Self-thinning to Artificially Reduced Levels of Leaf Area in Monocultures of Trifolium pratense

Greenhouse grown monocultures of Trifolium pratense L, werepartially defoliated to test the hypothesis that the constantof proportionality (K) in the self-thinning rule is a functionof leaf area. This constant equates mean weight (w) and density(p) in self-thinning populations by the equation Pruning leaflets did not affect the pattern or occurrence ofdensity-related mortality but, as predicted, did affect K, whichwas reduced by 25 per cent as leaf area was decreased from anaverage of 7.3 to 3.9 m²m^–2. For both self-thinning andnon-self-thinning populations, leaf area was substituted forK in eqn (1) to give Multiple linear regression showed that this expression was significantfor all three defoliation treatments. Regressions with treedata grouped by genera were also significant and indicate thateqn (2) may be a more general expression of the relation betweenmean weight and density in even-aged monocultures. The self-thinningrule may be a special case of eqn (2) which expresses itselfwhen leaf area attains some upper limit. Trifolium pratense, red clover, leaf area, self-thinning, defoliation     

Light Dependency of Carboxylation Efficiency and Ribulose-1, 5-Bisphosphate Carboxylase Activation in Trifolium subterraneum L. Leaves

The effect of photosynthetic photon flux density (PPFD) on carboxylationefficiency, estimated as the initial slope (IS) of net CO₂ assimilationrate versus intercellular CO₂ partial pressure response curve,as well as on ribulose-1, 5-bisphosphate carboxylase (Rubisco)activation was measured in Trifolium subterraneum L. leavesunder field conditions. The relationship between IS and PPFDfits a logarithmic curve. Rubisco activation accounts for theIS increase only up to a PPFD of 550 µmol photons m^-2s^-1. Further IS increase, between 550 and 1000 µmol photonsm^-2 s^-1, could be related to a higher ribulose fcwphosphate(RuBP) availability. The slow, but sustained IS increase above1000 µmol photons m^-2 s^-1 could be explained by the mesophyllCO₂ diffusion barriers associated with the high chlorophylland protein content in field developed leaves. Key words: Photosynthesis, initial slope, ribulose-1, 5-bissphosphate carboxylase activation, light response, Trifolium subterraneum L     

Male Sterility and Anther Wall Structure in Copper-deficient Plants

Anther development and pollen sterility were followed in plantsof wheat, oat, barley, sweetcorn, sunflower, petunia and subterraneumclover grown at a range of copper supplies. Copper-deficientplants had increased pollen sterility. Lignified wall thickenings were reduced or absent in the endotheciaof anthers from Cu-deficient plants. Reduced seed set may resultboth from reduced pollen fertility or failure of the stomiato rupture due to decreased lignification of anther walls. Triticum aestivum L., wheat, Hordeum vulgare L., barley, Avena sativa L., oat, Zea mays L., corn, sweetcorn, maize, Helianthus annuus L., sunflower, Petunia hybrida L., Trifolium subterraneum L., subterranean clover, male sterility, anther development, copper deficiency     

A Dynamic Equation for Plant Interaction and Application to Yield-density-time Relations

A model for plant interaction is developed based on a definitionof space in terms of actual and potential amount of growth factorsabsorbed per unit of time. The resulting equation is a second-orderdifferential equation which is solved by dynamic simulation.Five data sets on yield-density relations are used to demonstratethe model's excellent predictive power. Competition model, plant interaction, yield-density relations, Richards function, subterranean clover (Trifolium subterraneum L.), Wimmera ryegrass (Lolium loliaceum Hand-Mazz.), lettuce (Lactuca sativa L.), maize (Zea mays L.), lucerne (Medicago sativa L     

Callose Deposits make Clover Seeds Impermeable to Water

The permeability status of the seed from ten genotypes of subterraneanclover, determined by soaking, is shown to be related to callosedeposition in the seed coat. Large amounts of callose were foundin the nutrient (parenchyma) layer of impermeable but not permeableseeds. It is suggested that the nutrient layer controls permeabilityand thus the ease of germination of seeds. Trifolium subterraneum L., clover, seed coat impermeability, callose     

The Relationship Between the Phosphorus Status of Subterranean Clover Plants and Losses in Chlorophyll from Detached Leaves: A Basis for the Visual Diagnosis of Phosphorus Deficiency

Leaves detached from subterranean clover plants (Trifolium subterraneumL.), grown in solution cultures at different phosphorus levels,were placed in water and in phosphate solutions. Losses in totalchlorophyll (a+b) of leaves in water were greater the lowerthe previous phosphorus supply to the plants from which theywere detached. In comparable leaves placed in phosphate solutionschlorophyll was maintained at levels at least as high as inleaves from non-deficient plants. For the latter there wereno differences between treated and untreated leaves. The differencesin colour change between treated and untreated leaves, therefore,increased with the deficiency in the phosphorus supply to theplants from which the leaves were detached, thus visibly reflectingtheir phosphorus status. A light intensity of 300 ft c (27 lx) or higher was necessaryfor maximum differences in leaf colour between treated and untreatedleaves, mainly because lower light levels reduced chlorophylllosses from leaves in water. Differences in colour between treated and untreated deficientleaves decreased with their age. However, valid comparisonsbetween treated and untreated leaf tissue could be ensured bythe use of treated and untreated leaflets. It is suggested that the results provide the basis for a simple,direct and visual diagnostic method, requiring no laboratoryequipment. Trifolium subterraneum L., subterranean clover, phosphorus deficiency, chlorophyll     

Effects of Temperature on Growth and Nitrogen Fixation by Swards of Subterranean Clover

Subterranean clover (Trifolium subterraneum L.) cv. ‘Woogenellup’ swards were grown at 10, 15, 20 and 25 Cwith a 12 h photoperiod of 500 or 1000 µmol m^–2s^–1 [low and high photosynthetic photon flux density (PPFD)].Nitrogen-fixing swards received nutrient solution lacking combinednitrogen while control swards received a complete nutrient solution.Growth was measured by infra-red analysis of carbon dioxideexchange and by accumulation of dry matter. Swards were harvestedat intervals between 95 and 570 g d. wt m^–2 for estimationof nitrogenase activity by acetylene reduction and hydrogenevolution assays. Nitrogen fixation was also measured by increasein organic nitrogen. The growth rate was highest at 10 C at low PPFD, and at 10–15C at high PPFD. Nitrogen-fixing swards grew slower than thosereceiving combined nitrogen. Nitrogen fixation measured by increasein organic nitrogen responded similarly to the growth rate,as did acetylene reduction between 10 and 20 C. At 25 C therelationship between acetylene reduction and nitrogen fixationwas distrupted. The difference between the rates of acetylenereduction and hydrogen evolution, theoretically proportionalto nitrogen fixation, was not a reliable indicator of nitrogenfixation because hydrogen uptake developed. Trifolium subterraneum L, subterranean clover, growth, nitrogen fixation, temperature, acetylene reduction     

Modelling the Time Course of Self-thinning in Crowded Plant Populations

A logarithmic model for the self-thinning of plants is proposed.This model describes the time course of self-thinning very welland fits data from forest stands and yield tables, which followthe 3/2 power law. An approximated expression of this modelshows that plant density decreases with age along a Gompertzcurve. This appears to be a basic property of the time courseof self-thinning in plants. Pinus strobus L., Pinus densiflora Sieb, et Zucc., stand development, self-thinning, 3/2 power law, logarithmic model, mortality     

The Structure and Development of Trifolium subterraneum L. Root Nodules: I. IN PLANTS GROWN AT OPTIMAL ROOT TEMPERATURES

The structure of effective nodules on Trifolium subterraneumL. grown at a root temperature favourable for nitrogen fixationwas examined by light microscopy. Rhizobium were released intohost cells from vesicles on infection threads which were closelyassociated with the host cell nucleus. Enlargement into bacteroidforms was rapid and synchronous in similarly-aged host cells,but groups of rhizobia close to the nucleus sometimes did notdevelop into bacteroids. Rhizobium changed in shape from smallrods to more elongate and then to irregularly swollen formsbefore finally becoming coccoid. On degeneration clusters ofbacteroids became un evenly stained before coalescing at thecentre of the host cell and finally disintegrating. Small vegetativerods multiplied amongst the degenerating bacteroids and in thegreatly enlarged intercellular spaces; host cell walls becamefolded and distorted and sometimes broke before collapsing ontoeach other.     

Interrelations Between Seeds and Associated Fruiting Structures During Their Development in Trifolium subterraneum L.

Developing burrs of T. subterraneum (cv. Daliak) were subjectedto dark and light treatments in experiments conducted in a controlledtemperature glasshouse with day/night air temperatures of 18/13°C. Post fertilization development of burr components wasfollowed and the effect of removal of seeds or sterile calyoeson the development of the remaining burr components studied. Peduncle elongation took place mainly between days 0 (anthesis)and 7, when the burr would normally have buried. Seed developmentwas completed by day 42. Pod walls attained near maximum weightat about day 22. The perianth increasd in weight between days12 and 22 due to enlargement of the calyx. Sterile calyces wereprduced continuously until day 42. The pattern of burr develompentwas similar in both dark and light, but the weight of the maincomponents were considerably reduced in the light treatment. Normal peduncle development depended on the presence of at leastone seed in the inflorescence. Pod walls did not develop, neitherdid the perianth enlarge unless these structures were associatedwith a developing seed. In the absence of seeds fewer sterilecalyces were produced in the dark treatment and fewer stillwere produced in the light, indicating a direct inhibitory effectof light on the production of sterile calyces. Within matureburrs the weight of pod walls and sterile calyces was closelyrelated to the weight of seeds. Removal of sterile calyces reducedseed production but interpretation of this effect is not clear. Trifolium subterraneum L., fruit development, seed production     

Flowering Sequence and the Ordering of Impermeability Breakdown in Seed of Subterranean Clover (Trifolium subterraneum L.)

The pattern of breakdown of seed coat impermeability of seedwithin the burrs of two cultivars of subterranean clover (Trifoliumsubterraneum L.) was closely associated with relative differencesin flowering sequence within the inflorescence. The patternof seed weight variation showed very little relationship withthe pattern of breakdown, while that for seed moisture contentshowed no relationship. Within a burr it was almost invariably the seed which developedfrom the latest flower within the inflorescence which was thefirst to become permeable. An ordering of the timing of breakdownwas evident only within the burr; there was no evidence fora pattern of breakdown of impermeability between the differentburrs along the runner. Trifolium subterraneum L., subterranean clover, seed coat impermeability, seed weight, seed moisture, hardseededness     

Regrowth by Swards of Subterranean Clover after Defoliation. 1. Growth, Non-structural Carbohydrate and Nitrogen Content

Swards of subterranean clover (Trifolium subterraneum L.) atLAl 6 grown in N-free nutrient solution were subjected to threedefoliation treatments which removed 30, 70 and 80% of shootdry weight. Subsequent regrowth and changes in the concentrationsof carbohydrate and nitrogen in plant components were measuredat 0, 1, 5, 9 and 13 d after defoliation and compared with thosein uncut swards. The rate of shoot regrowth declined with increasing severilyof defoliation. In all defoliation treatments, growth was confinedto leaves for up to 5 d. Root growth ceased in all treatmentsfor a longer period. Reestablishment of the leaf area in severely-defoliatedswards was facilitated by the rapid opening of developing leavesand by changes in the allocation of carbon which favoured leafover branch and root, and lamina over petiole growth. Loss of carbohydrate and nitrogen from roots and branches lasting5–9 d was observed in the more severe defoliation treatments.Loss of protein (N x 6.25) exceeded that of total non-structuralcarbohydrate, and could have accounted for the nitrogen contentof new leaf during this period. Branches lost 62% of their initialcarbohydrate content compared with 25% from roots in the 80%cut swards. In contrast, roots, by virtue of their greater mass,were the principle source of mobilized nitrogen. Nitrogen accumulationceased in 80% cut swards for 9 d. However, carbohydrate levelsin the crown nodules were not severely depleted. It was concluded that partitioning of growth to lamina and themobilization of carbohydrates and nitrogen were important forrecovery from defoliation. Carbohydrates, carbon partitioning, defoliation, nitrogen, mobilization, regrowth, subterranean clover, Trifolium subterraneum L     

Physiological Predetermination of the Order of Hardseededness Breakdown in Subterranean Clover (Trifolium subterraneum L.)

Hardseededness breakdown in the burr of subterranean clover(Trifolium subterraneum L.) was found to be non-random in occurrencewith respect to the relative position of seed in the burr. Thissuggests the presence of a predeterminative mechanism of theorder of hardseededness breakdown. While seed moisture content did not exhibit a pattern of variationconsistent with that for hardseededness breakdown, seed weightvariation appeared to conform closely with the pattern of breakdown. It is postulated that, either differences in seed size per sewithin the burr may determine the order of hardseededness breakdown,or that such differences may reflect a disproportionate supplyof translocate, and hence promotors or inhibitors, of hardseedednessbreakdown to the seeds of the burr. The implications of this finding to the assessment of genotypesfor differences in level of hardseededness level have been outlined.     

Some Effects of Potassium and Magnesium on the Growth of Subterranean Clover (Trifolium subterraneum)

Subterranean clover plants (Trifolium subterraneum L., cv. Mt.Barker) were grown in culture solutions at optimum nutrientlevels and on the 23rd day after sowing transferred to solutionswithout magnesium or potassium. A third group remained at thesame nutrient levels as before (controls). Magnesium deficiency caused a particularly rapid decline inroot growth, followed later by a net loss of root dry matter.This, and the fact that leaf expansion declined relatively morethan the increase in laminae dry matter, resulted in an accumulationof assimilates per unit leaf area. Transfer of magnesium-deficientplants to complete solutions on day 35 caused a preferentialdistribution of dry matter to the roots, then to petioles andrelatively less to the laminae. These changes caused a pronouncedfall in the root: shoot ratio as the deficiency became moresevere and a rise in the ratio during recovery. Plants in solutions without potassium showed no marked shiftsin dry matter distribution between plant parts. The root: shootratio remained close to that for control plants, except duringthe recovery, when there was a decrease in the ratio. Net rates of CO₂ uptake by laminae from potassium-deficientplants showed little change during the first 10 days of thedeficiency although values were somewhat lower than those forcorresponding control laminae. After transfer to complete solutionsthere was a marked response in photosynthesis, rising to a finalvalue close to that for control laminae. Laminae of plants placedin solutions without magnesium showed a rapid decline in photosynthesisonly 4 days later; there was little response when plants weretransferred to complete solutions a week later. Trifolium subterraneum L., subterranean clover, growth, root: shoot ratio, potassium deficiency, magnesium deficiencies     

Growth and root dry matter allocation by pasture legumes and a grass with contrasting external critical phosphorus requirements

Background and aims

This work aimed to quantify the critical external requirement for phosphorus (P) (i.e. extractable-P concentration required for 90 % of maximum yield) for a number of temperate legume species and understand differences in dry matter allocation, P distribution and P acquisition efficiency among these species.

Methods

Shoot and root growth of five legume and one grass species was assessed in response to six rates of P mixed into the top 45 mm of soil in a pot experiment. Dactylis glomerata and Trifolium subterraneum were used as benchmark species; they are commonly grown together in mixed temperate pastures and have low and high critical external requirements for P, respectively. Growth was compared with four alternative legume species: Ornithopus compressus, Ornithopus sativus, Biserrula pelecinus and Trifolium hirtum, that have root morphologies better suited to soil exploration and nutrient acquisition than that of Trifolium subterraneum.

Results

Dactylis glomerata, Ornithopus compressus and Ornithopus sativus had maximum yields equal to or greater than Trifolium subterraneum but achieved this at rates of P less than half that of Trifolium subterraneum. Biserrula pelecinus and Trifolium hirtum had critical P requirements between that of Trifolium subterraneum and the Ornithopus species, but also had lower yields. Root dry matter of Dactylis glomerata and the Ornithopus species in the fertilised soil layer was only marginally changed in response to low P supply. In contrast, Trifolium subterraneum, Trifolium hirtum and to a lesser extent Biserrula pelecinus markedly increased root dry matter allocation to this soil layer. Species with lower critical P requirements were able to take up more P per unit root dry mass than those with higher critical P requirements, particularly at lower levels of P addition.

Conclusions

The high P acquisition efficiencies of the Ornithopus species and Dactylis glomerata were likely to have contributed to their low critical external P requirements. It was surmised that differences in root morphology traits underpin the differences in acclimation to low P stress and P acquisition efficiency among the species.

     

Assessment of Enzyme Induction and Aerenchyma Formation as Mechanisms for Flooding Tolerance in Trifolium subterraneum 'Park'

The objective of this study was to evaluate the role of enzymeinduction and aerenchyma formation in prolonged tolerance tosoil flooding in a variety of underground clover (Trifoliumsubterraneum ‘Park’) previously selected for resistance.Seedlings were grown in hydroponic tanks, initially with aerationfor 3 weeks and subsequently in the absence of aeration forup to 3 weeks. After 1 h in the absence of aeration, theoxygen concentration in the hydroponic medium had decreasedto 1·5 %. During the 3 weeks of extreme oxygen deficiency,primary roots died and were replaced by considerable numbersof adventitious roots. Activities of many glycolytic and fermentativeenzymes increased in adventitious roots. Excised adventitiousroots were capable of immediate induction of ethanol in theabsence of lactate production, in association with energy chargehigher than that in excised roots of aerobically maintainedcontrols. Energy charge was even higher when measured in adventitiousroots in planta. Interestingly, haemoglobin protein could becorrelated with energy charge. Aerenchyma was readily visualizedin adventitious roots by optical microscopy of longitudinaland transverse sections. We conclude that avoidance of rootanoxia via aerenchyma is the major mechanism for prolonged roottolerance in Trifolium subterraneum ‘Park’.     

A Canopy Photosynthesis Model for the Dynamics of Size Structure and Self-thinning in Plant Populations

A dynamic model for growth and mortality of individual plantsin a stand was developed, based on the process of canopy photosynthesis,and assuming an allometric relationship between plant heightand weight, i.e. allocation growth pattern of plant height andstem diameter. Functions G(t, x), for the mean growth rate ofindividuals of size x at time t, and M(t,x), for the mortalityrate of individuals of size x at time t, were developed fromthis model and used in simulations. The dynamics of size structurewere simulated, combining the continuity equation model, a simpleversion of the diffusion model, with these functions. Simulationsreproduced several well-documented phenomena: (1) size variabilityin terms of coefficient of variation and skewness of plant weightincreases at first with stand development and then stabilisesor decreases with an onset of intensive self-thinning; (2) duringthe course of self-thinning, there is a power relationship betweendensity and biomass per unit ground area, irrespective of theinitial density and of the allocation-growth pattern in termsof the allometric parameter relating plant height and weight.The following were further shown by simulation: (a) competitionbetween individuals in a crowded stand is never completely one-sidedbut always asymmetrically two-sided, even though competitionis only for light; (b) plants of ‘height-growth’type exhibit a greater asymmetry in competition than plantsof ‘diameter-growth’ type, (c) the effect of competitionon the growth of individuals in a crowded stand converges toa stationary state, even when the stand structure still changesgreatly. All of these theoretical results can explain recentempirical results obtained from several natural plant communities.Finally, a new, general functional form for G(t, x) in a crowdedstand is proposed based on these theoretical results, insteadof a priori or empirical growth and competition functions. Canopy photosynthesis, competition mode, continuity equation, self-thinning, simulation, size distribution