Analysis of Dry Matter Partitioning in Dactylis glomerata during Vegetative Growth Using a Carbon Budget Model

The dry matter partitioning in vegetative plants of Dactylisglomerata was studied from experiments performed in controlledenvironments. Plants were grown hydroponically in growth chambers,at two constant temperatures (17 and 25 °C). In both experimentsthe root fraction decreased regularly with time, an effect thatwas more accentuated in the higher temperature regime. In orderto explain the change in dry matter partitioning, the experimentalshoot and root growth were analysed using a carbon budget modelwhich includes shoot and root maintenance requirements. Themodel predicts a relationship between the root specific growthrate and the product of shoot specific growth rate and shootto root dry weight ratio. In the range of experimental accuracy,this relationship was found to be linear at both temperatures,which should indicate that the partitioning coefficients andthe root maintenance coefficient remained constant during vegetativegrowth. The effect of temperature on the value of these coefficientscan be specified from a linear regression analysis. Between17 and 25 °C, the root maintenance coefficient increasedby about a factor of two, whereas the partitioning coefficientsdid not vary significantly. On the basis of these results, itwas shown that the decrease in root fraction during vegetativegrowth should be mainly attributed to the decrease in net specificactivity of shoots.Copyright 1994, 1999 Academic Press Dactylis glomerata L., vegetative growth, model, partitioning, root:shoot ratio, shoot specific activity, maintenance requirements     

Effect of Nitrate Concentration on the Root: Shoot Ratio in Dactylis glomerata L. and on the Kinetics of Growth in the Vegetative Phase

The vegetative growth of Dactylis glomerata L. in sand was studiedunder controlled light, temperature, and nutritional conditions.Plants were daily supplied with three nutrient solutions ofdifferent nitrate concentrations (10^–2, 10^–3 and2 x 10^–4 mol I^–1). For each concentration, growthobeyed an exponential law between the fourth and seventh weeksafter sowing. The time constant of the exponential was the samefor the shoot as for the root, and showed no significant variationwith nitrate concentration. The kinetic results and the strong dependence of the root: shootratio on nitrate concentration are discussed on the basis ofThornley's model. Hypothesizing that the molecular mechanismsof nitrate absorption are independent of the nitrate concentrationof the nutrient solution, we derived a relationship betweenthe root: shoot ratio and nitrate concentration. This relationshipwas found to be compatible with the experimental results. Dactylis glomerata L., vegetative phase, kinetics of growth, root: shoot equilibrium, nitrate absorption     

Relative Nitrogen Limitation at Steady-state Nutrition as a Determinant of Plasticity in Five Grassland Plant Species

Partitioning of biomass between roots and different shoot partshas often been used to explain the response of plants to variationsin resource availability. There are still many uncertaintiesin the importance of this trait for plant performance, and clearguidelines on how partitioning should be quantified in relationto growth rate and resource supply are of fundamental importancefor such an understanding. This paper reports an attempt toshow how plant nitrogen status relates to root:shoot partitioningand other plastic responses, in a manner that can be used forquantitative predictions. The reactions to nitrogen limitationof five grassland plant species, with different ecological demands,were compared. The species used were the forbs Polygala vulgarisand Crepis praemorsa, and the grasses Danthonia decumbens, Agrostiscapillaris and Dactylis glomerata. The experiment was conductedin a climate chamber where the plants were grown hydroponically(1) under non-limiting nutrient conditions and (2) at a steady-statenitrogen limitation, which enabled the plants to express halfof their growth potential. The relative growth rate (RGR) ofthe species was strongly related to plant nitrogen concentration(PNC) and leaf area ratio (LAR), whereas the effects on netassimilation rate (NAR) were very small. Despite large differencesin maximum relative growth rate, the species showed remarkablesimilarities in dry matter partitioning between root and shoot.It is concluded that root:shoot partitioning can be treatedas a direct function of the relative resource limitation ofthe plant. The difficulty of attaining well-defined levels ofresource limitation in soil, other solid substrates and manyhydroponic systems may be the most important reason for thedivergent results in earlier studies. Better knowledge of soil-rootinteractions, and plant responses to the whole span of resource-supplylevels, is required for a thorough understanding of how nutrientslimit growth. Copyright 1999 Annals of Botany Company Growth rate, plant strategies, plasticity, partitioning, biomass, nitrogen, nutrient limitation, grassland.     

An Extension of the Logistic Model of Plant Growth

The kinetics of growth of Dactylis glomerata L. were studiedunder controlled temperature and nutritional conditions at threelevels of irradiance (35·55 and 85 W m^–s). Thedry weights of the root and shoot parts of the plants were measuredeach week between the fourth and eleventh weeks after sowing. The growth kinetics were found to be dependent on the levelof irradiance, but no significant differences in the root: totaldry weight ratio were observed. To characterize the effect of illuminance, the experimentalgrowth curves were analysed initially using the logistic model,the adequacy of which is discussed. An extension of the logisticmodel is proposed, represented by the kinetic equation dM/dt= kM, with < 1 and where M is the dry weight of the plant.It is shown that this relationship allows a distinction to bemade between two kinds of plant material according to theirfunctions in the growth process. Dactylis glomerata L., illuminance, growth curves, kinetic analyses, logistic model, shoot:root ratio, partition of assimilates     

Analysis of the Time Course of Change in Nitrogen Content in Dactylis glomerata L. Using a Model of Plant Growth

The growth, root fraction and nitrogen content of Dactylis glomerataL. grown hydroponically in constant environmental conditions,were measured in the vegetative phase. The variation of theseparameters denotes a progressive change in the chemical compositionof the plant, which can be attributed to changing proportionsof structural material, which is immediately available for growth,and storage material. On the basis of a kinetic model of plantgrowth, a definition of the two components was proposaed, whichled us to derive a relationship between relative growth rateand nitrogen content. This relationship was found to be compatiblewith experimental data. Dactylis glomerata L., vegetative phase, kinetic model, growth rate, nitrogen content     

Water Use Characteristics of Dactylis glomerata L., Lolium perenne L. and L. multiflorum Lam. Plants

Cocksfoot (Dactylis glomerata L.). perennial ryegrass (Loliumperenne L.) and Italian ryegrass (L. multiflorum Lam.) plantswere grown on deep (75–95 cm) columns of soil in glasshousesand growth rooms with and without irrigation. The species inwhich growth declined least rapidly after water had been withheldwere those which transpired most slowly. During early establishmentin the glasshouse cocksfoot transpired least because of slowroot growth. In the growth room, when root systems were deeperand denser, perennial ryegrass transpired least because of lowleaf water conductance. Results are discussed in relation to(a) drought resistance in the three species; (b) breeding forincreased drought resistance through modifying root distributionand leaf water conductance; and (c) the use of isolated soilcolumns in water relations studies. Dactylis glomerata L., Lolium perenne L., Lolium multiflorum Lam., cocksfoot, perennial ryegrass, Italian ryegrass, transpiration, roots, leaf water conductance     

Plant Growth Analysis: Allometry, Growth and Interference in Orchardgrass and Timothy

A multiple regression procedure was used to evaluate allometricresponses to stand age and species population densities in monoculturesand mixtures of orchardgrass (Dactylis glomerata L., also knownas cocksfoot) and timothy (Phleum pratense L.). In each speciesthe allometry between shoot dry weight and either leaf areaor tiller number per plant was studied. Population density treatmentsaffected allometry by changing allometric exponents expressingthe ratio of relative growth rates of different plant characteristics.Allometric relationships changed as growth proceeded, and thetwo species differed in their allometric responses to treatments. Plant growth analysis, allometry, competition, Dactylis glomerata L., Phleum pratense L.     

Nitrogen Uptake and Plant Growth II. An Empirical Model of Vegetative Growth and Partitioning

An empirical model of vegetative plant growth is presented.The model is based on experimental data on Polygonum cuspidatum,which showed (1) that the partitioning of dry matter and nitrogenamong organs was linearly related to the nitrogen concentrationof the whole plant and (2) that leaf thickness was negativelycorrelated with leaf nitrogen concentration. The model properlydescribes the behaviour of plants. Steady-state solutions ofthe model give the relative growth rate, specific leaf weight,and partitioning of dry matter and nitrogen among organs withthe net assimilation rate and the specific absorption rate asenvironmental variables. The effect of nitrogen removal on drymatter and nitrogen partitioning was examined as non-steady-statedynamic solutions of the model. The model predicted not onlyreduced leaf growth and enhanced root growth but also a fluxof nitrogen from the leaf to the root, which agreed with theexperimental results. Mathematical model, partitioning of dry matter and nitrogen, plant nitrogen, relative growth rate, shoot: root ratio, specific leaf weight     

Dry matter allocation and nitrogen productivity explain growth responses to photoperiod and temperature in forage grasses

The mechanisms responsible for fluctuations in species composition of semi-natural grassland are not well understood. To identify plant traits that determine the poor competitive ability of Festuca pratensis compared to Dactylis glomerata especially during summer, the growth of both grasses was monitored over time and at different temperatures and photoperiods. Plants of both grasses were grown from seed with non-limiting nutrient supply at three day/night temperatures (11/6, 18/13 and 25/20°C) and two photoperiods (16 and 12 h). F. pratensis had a significantly lower relative growth rate than D. glomerata, mainly due to its lower specific leaf area and reduced nitrogen productivity. At high temperature, F. pratensis had a considerably lower root weight ratio than D. glomerata leading to substantially slower root growth. F. pratensis responded to a shorter photoperiod with an increase in the net assimilation rate, whereas D. glomerata responded with an increase in specific leaf area. The low competitive ability of F. pratensis compared to D. glomerata was mainly associated with its lower specific leaf area and nitrogen productivity. The stronger decline of its competitive ability during summer was probably related to the decreased allocation of dry matter to the roots at higher temperatures which leads to slower root growth compared to D. glomerata. Received: 7 September 1998 / Accepted: 29 July 1999     

Leaf Extension Rate in Temperate Pasture Grasses in Relation to Assimilate Pool in the Extension Zone

In two growth cabinet experiments the leaf extension rate (LER)was studied under a 14 h photoperiod followed by prolonged darkness,in tillers of the perennial temperate pasture grasses Phalaristuberosa L. cv. Sirosa and Dactylis glomerata L. cv. Currie.Levels of soluble non-structural carbohydrates and total     

Is Partitioning of Dry Weight and Leaf Area Within Dactylis glomerata Affected by N and CO2Enrichment?

We examined changes in dry weight and leaf area within Dactylisglomerata L. plants using allometric analysis to determine whetherobserved patterns were truly affected by [CO₂] and N supplyor merely reflect ontogenetic drift. Plants were grown hydroponicallyat four concentrations of in controlled environment cabinets at ambient (360 µll^–1) or elevated (680 µl l^–1) atmospheric[CO₂]. Both CO₂and N enrichment stimulated net dry matter production.Allometric analyses revealed that [CO₂] did not affect partitioningof dry matter between shoot and root at high N supply. However,at low N supply there was a transient increase in dry matterpartitioning into the shoot at elevated compared to ambient[CO₂] during early stages of growth, which is inconsistent withpredictions based on optimal partitioning theory. In contrast,dry matter partitioning was affected by N supply throughoutontogeny, such that at low N supply dry matter was preferentiallyallocated to roots, which is in agreement with optimal partitioningtheory. Independent of N supply, atmospheric CO₂enrichment resultedin a reduction in leaf area ratio (LAR), solely due to a decreasein specific leaf area (SLA), when plants of the same age werecompared. However, [CO₂] did not affect allometric coefficientsrelating dry weight and leaf area, and effects of elevated [CO₂]on LAR and SLA were the result of an early, transient stimulationof whole plant and leaf dry weight, compared to leaf area production.We conclude that elevated [CO₂], in contrast to N supply, changesallocation patterns only transiently during early stages ofgrowth, if at all. Copyright 2000 Annals of Botany Company Allometric growth, carbon dioxide enrichment, Cocksfoot, Dactylis glomerata L., dry weight partitioning, leaf area ratio, nitrogen supply, shoot:root ratio, specific leaf area     

Effects of Soil Temperature on the Growth and Hormone Content of Dactylis glomerata L. (Cocksfoot) in Controlled Environments

A comparison was made of effects of ‘low’ and ‘high’soil temperature (LST and HST, about 9 and 21·5 °Crespectively) on shoot growth of Norwegian and Portuguese populationsof Dactylis glomerata. In experiments lasting 8 short days (SD,8 h photoperiods) LST decreased leaf extension more markedlyin the Portuguese population. No differential effect of LSTon leaf growth was recorded in experiments lasting 20 or 21SD or in experiments of 8 d duration in long days (LD, 16 hphotoperiods). Since the meristem and region of cell extensionis close to the soil surface LST could directly influence bothroot and shoot growth. The application of gibberellic acid enhanced leaf extension,particularly in plants grown at HST. 6-Benzylaminopurine tendedto decrease leaf length. Extractable gibberellin levels wererelatively low after 8 SD in shoots of both races grown at LST.Cytokinin levels increased at LST, more so in shoots of Portugueseplants which sustained the greater reduction in leaf extension. The data suggest that LST may reduce the production of endogenousgibberellins important for leaf growth in Dactylis glomerata.Cytokinins are probably necessary for growth processes but theirlevel may reflect, rather than direct, the rate of leaf growth.     

The Effect of Temperature on Vegetative Growth in Climatic Races of Dactylis glomerata in Controlled Environments

The growth patterns in two natural populations of Dactylis glomeratafrom contrasting climatic regions, Norway and Portugal, werestudied at four constant temperatures (5, 10, 20, and 30°C) in a 16-h photoperiod. Marked changes in relative growth-rateat different temperatures were positively correlated with changesin both net assimilation rate and leaf-area ratio, whereas differencesbetween the populations in the relative growth-rate were theresult of differences in net assimilation rate, and were negativelycorrelated with differences in leaf-area ratio. The changesin leaf-area ratio at different temperatures were correlatedwith changes in leaf morphology and distribution of assimilateswithin the plant. The possible adaptive advantage of these vegetativegrowth patterns is discussed in relation to the survival ofthe plants in the original environments.     

The Effect of Sulphur Dioxide on the Growth of Lolium perenne L., Lolium multiflorum Lam., Dactylis glomerata L., and Phleum pratense L.

Lockyer, D. R. 1985. The effect of sulphur dioxide on the growthof Lolium perenne L., Lolium multiflorum Lam., Dactylis glomerataL., and Phleum pratense L.?J. exp. Bot. 36: 1851-1859. Fouragriculturally important grasses, Lolium perenne L., Loliummultiflorum Lam., Dactylis glomerata L. and Phleum pratenseL. were exposed to sulphur dioxide (SO₂) in a system of exposurechambers. The plants were exposed for a total of 43 d to meanconcentrations of SO₂ in the air of 0,87 or 448 (µg m^–3and herbage was harvested twice. All four grasses showed chloroticlesions after exposure to the highest concentration of SO₂.The effect of SO₂ on the yield of herbage was statisticallysignificant only at the second harvest and at the highest concentration;the dry weights of shoots of D. glomerata and L. perenne werereduced by 33% and 16% respectively. Significant effects ofSO₂ were also found on the 'transpiration coefficients' measuredfor D. glomerata and P. pratense. The grasses differed in theiruptake of sulphur from the atmosphere but this was not relatedto their sensitivity to SO₂. Total–S concentration inthe shoots of L. perenne, L. multiflorum and D. glomerata increasedalmost linearly in response to increasing SO₂ concentration;with P. pratense only the highest SO₂ concentration raised total-Sabove the level in control plants. These increases were almostentirely due to the accumulation of sulphate–S. Key words: Sulphur diozide, Lolium perenne, Lolium multiflorum, Dactylis glomerata, Phleum pratense     

Effects of Drought on Water Relations, Mineral Uptake, Water-soluble Carbohydrate Accumulation and Survival of Two Contrasting Populations of Cocksfoot (Dactylis glomerata L.)

The role physiological responses in survival of prolonged soilmoisture deficit was investigated in vegetative plants of twoDactylis glomerata populations: KM2 (drought-resistant) andLutetia (susceptible). The plants were grown in 1 m-deep soilcolumns in a controlled environment. After 56 d of full irrigation,water was withheld for 80 d, by which time all soil moisturehad been consumed; the plants were then rewatered for a further19 d. As drought progressed, leaf extension decreased to zero,water status declined, and water-soluble carbohydrates (WSC)at first increased and then decreased. The most pronounced differencesbetween the two populations was that all KM2 tillers survivedthe drought, but 34% of Lutetia tillers died. In comparisonwith Lutetia, KM2 was characterized by (a) slower shoot growthrate, (b) greater root density at depth, (c) maintenance ofhigher lamina relative water content, (d) greater osmotic adjustmentin leaf bases, (e) higher concentration of WSC in tiller bases,(f) greater ability to export WSC out of dying leaves, (g) lowercontent of metal ions but improved maintenance of P status,and (h) lower proline:amino acid ratio. The contribution ofthese responses to tiller survival under severe drought in controlledenvironments is contrasted with performance and persistenceof swards in the field in the harsher Mediterranean environment.Copyright1995, 1999 Academic Press Dactylis glomerata L., cocksfoot, water-stress, osmotic asjustment, drought survival, root-growth, water-soluble carbohydrates, phosphate, proline, variety comparison     

Root and leaf attributes accounting for the performance of fast- and slow-growing grasses at different nutrient supply

Despite their difference in potential growth rate, the slow-growing Brachypodium pinnatum and the fast-growing Dactylis glomerata co-occur in many nutrient-poor calcareous grasslands. They are known to respond differently to increasing levels of N and P. An experiment was designed to measure which characteristics are affected by nutrient supply and contribute to the ecological performance of these species. Nutrient acquisition and root and shoot traits of these grasses were studied in a garden experiment with nine nutrient treatments in a factorial design of 3 N and 3 P levels each. D. glomerata was superior to B. pinnatum in nutrient acquisition and growth in all treatments. B. pinnatum was especially poor in P acquisition. Both species responded to increasing N supply and to a lesser extent to increasing P supply by decreasing their root length and increasing their leaf area per total plant weight. D. glomerata showed a higher plasticity. In most treatments, the root length ratio (RLR) and the leaf area ratio (LAR) were higher for D. glomerata. A factorization of these parameters into components expressing biomass allocation, form (root fineness or leaf thickness) and density (dry matter content) shows that the low density of the biomass of D. glomerata was the main cause for the higher RLR and LAR. The biomass allocation to the roots showed a considerable plasticity but did not differ between the species. B. pinnatum had the highest leaf weight ratio. Root fineness was highly plastic in D. glomerata, the difference with B. pinnatum being mainly due to the thick roots of D. glomerata at high nutrient supply. The leaf area/leaf fresh weight ratio did not show any plasticity and was slightly higher for B. pinnatum. It is concluded, that the low density of the biomass of D. glomerata is the pivotal trait responsible for its faster growth at all nutrient levels. It enables simultaneously a good nutrient acquisition capacity by the roots as well as a superior carbon acquisition by the leaves. The high biomass density of B. pinnatum will then result in a lower nutrient requirement due to a slower turnover, which in the long term is advantageous under nutrient-poor conditions.     

The Role of Different Daily Irradiations on Shoot Growth and Root/Shoot Ratio in Lucerne (Medicago sativa L.)

The effect of two levels of daily irradiation on shoot growthand root/shoot partitioning was investigated on two lucernevarieties (Medicago sativa L.). Individual plants were studiedunder constant temperature and optimal water and mineral nutritionconditions. For both lucerne varieties, daily irradiation did not changebiomass partitioning between shoots and roots. It can be shownthat leaf area expansion occurs independently of daily irradiationand that the process of shoot dry matter production is strictlyproportional to daily irradiation, since the ratio leaf area/shootdry matter is inversely proportional to it. From a model of shoot production of the isolated plant, we showthat the relative shoot specific activity is not affected bydaily irradiation. A simple model of partitioning leads us todetermine why partitioning remains the same for the two at thelevels of daily irradiation. Finally, the allocation between shoot and root turns out tobe independent of the main stem extension rate, which is fasterat the higher daily irradiation. Medicago sativa L., lucerne, daily irradiation, shoot production, root/shoot ratio, leaf area expansion, partitioning model