Growth response to step-decrease in nutrient availability in small birch (Betula pendula Roth)

Abstract Changes in the uptake and allocation of carbon and nitrogen, after a step-decrease in nutrient availability, were investigated in small birch (Betula pendula Roth). By demonstrating stable nutrition, before and after the decrease in nutrient supply, it was possible to eliminate the effects of plant size and age. Immediately following the step-decrease in nutrient availability, net nitrogen uptake to leaves and the relative rate of increase in shoot area tended to zero. Although photosynthetic rate per shoot area decreased, carbon uptake remained in excess of that used in structural growth and respiration. More of the excess carbon was accumulated as starch in leaves than in roots. After a lag phase, the relative rates of increase in plant dry matter, starch amount, net nitrogen uptake to leaves and shoot area development equalled that of the reduced rate of nutrient supply. It is concluded that the reduction in plant relative growth rate was much more attributable to the reduced allocation of photosynthate to leaf area growth than to the reduction in photosynthesis per shoot area.     

A dynamic model for nitrogen-stressed lettuce

A previously developed dynamic lettuce model, designed to predict growth and nitrate content under the normal range of glasshouse environmental conditions, has been extended to cover high nitrogen-stress situations. Under severe shortage of nitrogen, lettuce has been observed to grow at a very slow rate, as well as to have abnormally low water content, low reduced-nitrogen content and negligible nitrate content. The new model mimics these observations by adding to the original model a storage compartment for 'excess' carbon. The resulting model has three compartments: (1) 'vacuole', where the soluble non-structural material is stored, and the nitrate : carbon ratio may vary as needed to maintain a constant osmotic potential; (2) 'structure', a metabolically active compartment with fixed chemical composition; and (3) 'excess-carbon', which serves as a long-term storage of 'waterless' carbohydrates. Simulations with the model illustrate its ability to predict the effect of light, temperature and nitrogen in the nutrient solution on the long-term growth and composition of lettuce. They also illustrate the effects of plant size, and the associated relative growth rate, on the characteristic times of transient responses resulting from step changes in the environment.     

Nutrient and water availability alter belowground patterns of biomass allocation, carbon partitioning, and ectomycorrhizal abundance in Betula nigra

In managed settings, seedlings are often fertilized with the objective of enhancing establishment, growth, and survival. However, responses of seedlings to fertilization can increase their susceptibility to abiotic stresses such as drought. Seedlings acclimate to variation in soil resources by reallocating carbon among different physiological processes and compartments, such as above versus belowground growth, secondary metabolism, and support of ectomycorrhizal fungi (EMF). We examined the effects of nutrient and water availability on carbon allocation to above and belowground growth of river birch (Betula nigra), as well as partitioning among root sugars, starch, phenolics, lignin, and EMF abundance. As nutrient availability increased, total plant biomass and total leaf area increased, while percent root biomass decreased. Root sugars, total root phenolics and EMF abundance responded quadratically to nutrient availability, being lowest at intermediate fertility levels. Decreased water availability reduced total leaf area and root phenolics relative to well-watered controls. No interactions between nutrient and water availability treatments were detected, which may have been due to the moderate degree of drought stress imposed in the low water treatment. Our results indicate that nutrient and water availability significantly alter patterns of carbon allocation and partitioning in roots of Betula nigra seedlings. The potential effects of these responses on stress tolerance are discussed.     

根茎型植物扁秆荆三棱对光照强度和养分水平的生长响应及资源分配策略

自然界中光照和养分因子常存在时空变化,对植物造成选择压力。克隆植物可通过克隆生长和生物量分配的可塑性来适应环境变化。尽管一些研究关注了克隆植物对光照和养分因子的生长响应,但尚未深入全面了解克隆植物对光照和养分资源投资的分配策略。以根茎型草本克隆植物扁秆荆三棱(Bolboschoenus planiculmis)为研究对象,在温室实验中,将其独立分株种植于由2种光照强度(光照和遮阴)和4种养分水平(对照、低养分、中养分和高养分)交叉组成的8种处理组合中,研究了光照和养分对其生长繁殖及资源贮存策略的影响。结果表明,扁秆荆三棱的生长、无性繁殖及资源贮存性状均受到光照强度的显著影响,在遮阴条件下各生长繁殖性状指标被抑制。且构件的数目、长度等特征对养分差异的可塑性响应先于其生物量积累特征。在光照条件下,高养分处理的总生物量、叶片数、总根茎分株数、长根茎分株数、总根茎长、芽长度、芽数量等指标大于其他养分处理,而在遮阴条件下,其在不同养分处理间无显著差异,表明光照条件可影响养分对扁秆荆三棱可塑性的作用,且高营养水平不能补偿由于光照不足而导致的生长能力下降。光照强度显著影响了总根茎、总球茎及大、中、小球茎的生物量分配,遮阴条件下,总生物量减少了对地下部分根茎和球茎的分配,并将有限的生物量优先分配给小球茎。总根茎的生物量分配未对养分发生可塑性反应,而随着养分增加,总球茎分配下降,说明在养分受限的环境中球茎的贮存功能可缓冲资源缺乏对植物生长的影响。在相同条件下,根茎生物量对长根茎的分配显著大于短根茎,以保持较高的繁殖能力;而总球茎对有分株球茎的生物量分配小于无分株球茎,表明扁秆荆三棱总球茎对贮存功能的分配优先于繁殖功能。研究为进一步理解根茎型克隆植物对光强及基质养分环境变化的生态适应提供了依据。     

Bacterial growth physiology and RNA metabolism

Bacteria are sophisticated systems with high capacity and flexibility to adapt to various environmental conditions. Each prokaryote however possesses a defined metabolic network, which sets its overall metabolic capacity, and therefore the maximal growth rate that can be reached. To achieve optimal growth, bacteria adopt various molecular strategies to optimally adjust gene expression and optimize resource allocation according to the nutrient availability. The resulting physiological changes are often accompanied by changes in the growth rate, and by global regulation of gene expression. The growth-rate-dependent variation of the abundances in the cellular machineries, together with condition-specific regulatory mechanisms, affect RNA metabolism and fate and pose a challenge for rational gene expression reengineering of synthetic circuits.This article is part of a Special Issue entitled: RNA and gene control in bacteria, edited by Dr. M. Guillier and F. Repoila.     

Predicting microbial growth dynamics in response to nutrient availability

Developing mathematical models to accurately predict microbial growth dynamics remains a key challenge in ecology, evolution, biotechnology, and public health. To reproduce and grow, microbes need to take up essential nutrients from the environment, and mathematical models classically assume that the nutrient uptake rate is a saturating function of the nutrient concentration. In nature, microbes experience different levels of nutrient availability at all environmental scales, yet parameters shaping the nutrient uptake function are commonly estimated for a single initial nutrient concentration. This hampers the models from accurately capturing microbial dynamics when the environmental conditions change. To address this problem, we conduct growth experiments for a range of micro-organisms, including human fungal pathogens, baker’s yeast, and common coliform bacteria, and uncover the following patterns. We observed that the maximal nutrient uptake rate and biomass yield were both decreasing functions of initial nutrient concentration. While a functional form for the relationship between biomass yield and initial nutrient concentration has been previously derived from first metabolic principles, here we also derive the form of the relationship between maximal nutrient uptake rate and initial nutrient concentration. Incorporating these two functions into a model of microbial growth allows for variable growth parameters and enables us to substantially improve predictions for microbial dynamics in a range of initial nutrient concentrations, compared to keeping growth parameters fixed.     

Plasticity of seed output in response to soil nutrients and density in Abutilon theophrasti: implications for maintenance of genetic variation

 Seed output is determined by two processes: resource acquisition and the allocation of resources to seeds. In order to clarify how the reaction norm of seed output is controlled by the phenotypic expression of its two components, we examined the genetic components of plasticity of seed dry mass, plant size, and reproductive allocation under different conditions of soil nutrient availability and conspecific competition among eight families of Abutilon theophrasti. Without competition, the reaction norm of seed mass of the families crossed between the lowest and other nutrient levels, although neither of its components, plant size and reproductive allocation, showed such a response. The crossing reaction norm (i.e., reversal of relative fitnesses of different genotypes along the environmental gradient) of seed mass resulted from (1) a trade-off between plant size and reproductive allocation, and (2) changes in the relative magnitude of genetic variances in plant size and reproductive allocation with soil nutrient availability. While allocation was more important in determining seed mass under limiting nutrient conditions, plant size became more important under high-nutrient conditions. There were no significant genetic variances in seed mass, plant size, and reproductive allocation in the competition treatment, except at the highest nutrient level. The results show that plant competition mitigated the effects of genetic differences in plant performance among the families. We discuss the results in relation to maintenance of genetic variation within a population. Received: 16 August 1996 / Accepted 26 April 1997     

Equilibrium and balanced growth of a vegetative crop

MODEL: A previously developed dynamic model, NICOLET, designed to predict growth and nitrate content of a lettuce crop, is subjected to (virtual) constant environmental conditions. For every combination of shoot and root environment, the cell sap, here assumed to reside in the "vacuole" compartment, equilibrates at a certain nitrate concentration level. This, in turn, defines the composition of the crop in terms of carbon and nitrogen content in each of the three compartments of the model. Growth under constant environmental conditions is defined as "equilibrium" growth (EG). If, in addition, the source strengths of carbon and nitrogen balance each other, as well as the sink strength of the growing crop, the growth is said to be "balanced" (BG). RESULTS: It is shown that the range of BG approximately coincides with the range of "mild" nitrogen stress, where reduction in nitrogen availability results in a mild reduction of relative growth rate (RGR). Beyond a certain low nitrate concentration in the cell sap, the N-stress becomes "severe" and the loss of growth increases considerably. CONCLUSIONS: The model is able to mimic the five central observations of many constant-environment growth-chamber experiments, namely (1) the initial exponential growth and later decline of the RGR, (2) the constant chemical composition, (3) the equality of the RGR and the relative nutrient supply rate (RNR), (4) the proportionality between the N : C ratio and the RNR, and (5) the proportionality between the water content and the reduced N content. Guidelines for the optimal combination of the shoot and root environments are suggested.     

An improved model of root growth in structured soil

Summary The growth of roots of maize, sorghum and soybean is modelled through beds of spherical aggregates. Effects of aggregate size and strength, and effects of the spread or distribution of aggregate strengths are investigated.This is achieved by a combination of a statistical model for soil structure with a statistical model for the penetration behaviour of a root at a void/aggregate interface. It is shown that the behaviour of a root at such an interfac is dependent on the previous history of the root in its passage through the soil.It is concluded that the smaller the aggregate size, the greater is the nutrient availability per unit length of root. The influence of aggregate size decreases with increasing soil strength.An increase in aggregate strength reduces the availability of nutrients per unit length of root. However, the rate of nutrient uptake per root axis goes through a minimum at a strength (for maize) of around 80 per cent of the maximum limiting aggregate strength for root penetration. An increase in the spread of aggregate strengths usually results in a proportional increase in nutrient availability. This effect is more pronounced with smaller aggregate sizes.     

The reallocation of carbon in P deficient lupins affects biological nitrogen fixation

It is not known how phosphate (P) deficiency affects the allocation of carbon (C) to biological nitrogen fixation (BNF) in legumes. The alteration of the respiratory and photosynthetic C costs of BNF was investigated under P deficiency. Although BNF can impose considerable sink stimulation on host respiratory and photosynthetic C, it is not known how the change in the C and energy allocation during P deficiency may affect BNF. Nodulated Lupinus luteus plants were grown in sand culture, using a modified Long Ashton nutrient solution containing no nitrogen (N) for ca. four weeks, after which one set was exposed to a P-deficient nutrient medium, while the other set continued growing on a P-sufficient nutrient medium. Phosphorus stress was measured at 20 days after onset of P-starvation. During P stress the decline in nodular P levels was associated with lower BNF and nodule growth. There was also a shift in the balance of photosynthetic and respiratory C toward a loss of C during P stress. Below-ground respiration declined under limiting P conditions. However, during this decline there was also a shift in the proportion of respiratory energy from maintenance toward growth respiration. Under P stress, there was an increased allocation of C toward root growth, thereby decreasing the amount of C available for maintenance respiration. It is therefore possible that the decline in BNF under P deficiency may be due to this change in resource allocation away from respiration associated with direct nutrient uptake, but rather toward a long term nutrient acquisition strategy of increased root growth.     

Cell size control in bacteria

Like eukaryotes, bacteria must coordinate division with growth to ensure cells are the appropriate size for a given environmental condition or developmental fate. As single-celled organisms, nutrient availability is one of the strongest influences on bacterial cell size. Classic physiological experiments conducted over four decades ago first demonstrated that cell size is directly correlated with nutrient source and growth rate in the Gram-negative bacterium Salmonella typhimurium. This observation subsequently served as the basis for studies revealing a role for cell size in cell cycle progression in a closely related organism, Escherichia coli. More recently, the development of powerful genetic, molecular, and imaging tools has allowed us to identify and characterize the nutrient-dependent pathway responsible for coordinating cell division and cell size with growth rate in the Gram-positive model organism Bacillus subtilis. Here, we discuss the role of cell size in bacterial growth and development and propose a broadly applicable model for cell size control in this important and highly divergent domain of life.     

Seed germination and seedling performance of two Mediterranean tree species,holm oak (shape Quercus ilex L.) and Aleppo pine (shape Pinus halepensis Mill.): a multifactor experimental approach

A two-level multifactor experimental approach was used to compare seed germination and seedling performance of two Mediterranean tree species: the early successional Aleppo pine (Pinus halepensis Mill.) and the late successional holm oak (Quercus ilex L.). In a first experiment germination rate was evaluated under the combined effects of shade, nitrogen availability, and pine or holm oak leaf litter. In a second experiment we tested for the effects of shade, nutrient availability, and litter type on seedling survival, growth and biomass allocation. Holm oak showed higher germination rates under shaded than under unshaded conditions, while Aleppo pine showed no differences between shaded and unshaded conditions. Nitrogen availability and litter type had no significant effect on germination of either species. Both species showed increased RGR, but also higher mortality rates, when grown in an enriched nutrient environment. While Aleppo pine showed no differences in RGR and mortality rate under different shading levels, RGR decreased and mortality increased for holm oak in full light. Increased radiation decreased LAR, SLA and height:diameter ratio, and increased RWR and R/S in both species, although Aleppo pine showed more pronounced changes. Unlike Aleppo pine, holm oak responded to increased nutrient availability by decreasing R/S and increasing LAR. From these results, no seed-seedling conflicts were found in either species, but a trade-off does seem to exist for holm oak between biomass allocation traits deployed in response to increased nutrient availability and radiation. Aleppo pine outperformed holm oak under most environmental conditions tested and showed a wider regeneration niche.     

Ecoenzymatic stoichiometry of microbial nutrient acquisition in tropical soils

The relative activities of soil enzymes involved in mineralizing organic carbon (C), nitrogen (N), and phosphorus (P) reveal stoichiometric and energetic constraints on microbial biomass growth. Although tropical forests and grasslands are a major component of the global C cycle, the effects of soil nutrient availability on microbial activity and C dynamics in these ecosystems are poorly understood. To explore potential microbial nutrient limitation in relation to enzyme allocation in low latitude ecosystems, we performed a meta-analysis of acid/alkaline phosphatase (AP), β-1,4-glucosidase (BG), and β-1,4-N-acetyl-glucosaminidase (NAG) activities in tropical soils. We found that BG:AP and NAG:AP ratios in tropical soils are significantly lower than those of temperate ecosystems overall. The lowest BG:AP and NAG:AP ratios were associated with old or acid soils, consistent with greater biological phosphorus demand relative to P availability. Additionally, correlations between enzyme activities and mean annual temperature and precipitation suggest some climatic regulation of microbial enzyme allocation in tropical soils. We used the results of our analysis in conjunction with previously published data on soil and biomass C:N:P stoichiometry to parameterize a biogeochemical equilibrium model that relates microbial growth efficiency to extracellular enzyme activity. The model predicts low microbial growth efficiencies in P-limited soils, indicating that P availability may influence C cycling in the highly weathered soils that underlie many tropical ecosystems. Therefore, we suggest that P availability be included in models that simulate microbial enzyme allocation, biomass growth, and C mineralization.     

Plasticity and ontogenetic drift of biomass allocation in response to above- and below-ground resource availabilities in perennial herbs: a case study of <Emphasis Type="Italic">Alternanthera philoxeroides</Emphasis>

Changes in plant biomass allocation in response to varying resource availabilities may result from ontogenetic drift caused by allometric growth (i.e., apparent plasticity), a true adjustment of ontogenetic trajectories (true plasticity) or both (complex plasticity). Given that the root allocation of annual species usually decreases during the growth, the developmentally explicit model predicts that annual herbs will exhibit true plasticity in root allocation under above-ground resource limitation and apparent plasticity for moderate stress of below-ground resource. For perennial species, the root allocation of which increases during growth, the reverse patterns would be expected. In this study, we tested the developmentally explicit model with a perennial weed, Alternanthera philoxeroides (Mart.) Griseb. We report its adaptive changes and ontogenetic drift of root allocation in response to different resource levels (i.e., light, water and nutrient availability) by comparing root allocation on both an age and a size basis. The root allocation of A. philoxeroides increased with the size (i.e., ontogenetic drift) during the growth, and exhibited significant changes in response to different resource availabilities. Furthermore, the root allocation in response to water or nutrient availability exhibited typical complex plasticity, while the light stress only slowed down the growth, with the ontogenetic trajectory unchanged (apparent plasticity). The contrasting responses to above-ground and below-ground stresses were consistent with the prediction of the developmentally explicit model.     

A Coordination Model of Whole-plant Carbon Allocation in Relation to Water Stress

Although water is an important determinant of the allocationof material between roots and shoots during growth, and oftenparallels the effects of nitrogen, few models have explicitlyconsidered allocation in relation to water supply. We use coordinationtheory to develop a simple exponential model that considersallocation of dry matter between shoots and roots during growthin relation to carbon and watersupplies, and accounts for theeffects of water stress on growth. We compare coordinationvs.optimization(global and local) versions of the exponential model by examiningsimilarities and differences in model behaviour obtained underconstant and variable environmental conditions, and with drasticallychanging conditions (mild, moderate and severe water stress).The greatest differences between coordination and optimizationexist in the drastically changing conditions. In a second versionof the model, we remove the restriction of exponential growthand show how coordination principles can be extended to a morecomplicated structure. The non-exponential model is used toanalyse experimental data on the effects of different pot sizes(and hence water availability) on root restriction and plantgrowth as reported by Thomas and Strain (Plant Physiology96:627–634, 1991). With further refinements, the coordinationmodel has potential as a tool to model plant growth in relationto water supply under various environmental conditions. Optimization; functional balance; root: shoot ratio; root restriction     

A conceptual model for the development of Phytophthora disease in Quercus robur

Here, a conceptual model is presented for the development of Phytophthora disease in pedunculate oak. The model is presented using the causal loop diagram tool and gives an overview of how various abiotic and biotic factors, such as soil moisture, nutrient availability and mycorrhizal colonization, may affect the reproduction and the infective capacity of soil-borne Phytophthora species, the susceptibility of the host and subsequent disease development. It is suggested that the link between the root damage caused by Phytophthora species and overall tree vitality is in the assimilation and allocation of carbon within the plants. The potential impact of environmental factors on these processes is discussed. The model is presented with reference to scenarios related to variation in soil moisture and nutrient availability. The need for species-specific validation of the model and the implications of the model are discussed.     

Effects of nutrient availability on biomass allocation as well as constitutive and rapid induced herbivore resistance in poplar

Many studies have examined effects of nutrient availability on constitutive herbivore resistance of plants, but few have addressed effects on expression of rapid induced resistance (RIR). We quantified effects of two levels of nutrient availability on growth, biomass allocation, photosynthesis, and constitutive secondary metabolism of black poplar (Populus nigra). We also examined effects of nutrient availability on expression of constitutive resistance of poplar to gypsy moth (Lymantria dispar) and whitemarked tussock moth (Orgyia leucostigma), as well as RIR to both folivores in response to localized herbivory by gypsy moth. The high nutrient treatment had no effect on photosynthetic rate of poplar, but dramatically increased relative growth rate, total biomass, and total leaf area, while foliar phenolic concentrations and root:shoot ratio decreased. Plant growth was negatively correlated with foliar phenolic concentrations, which is consistent with predictions of the Growth/Differentiation Balance Hypothesis when increased nutrient availability increases growth without affecting photosynthesis. These responses of root:shoot ratio and constitutive secondary metabolism to nutrient availability are consistent with those proposed by models of adaptive phenotypic plasticity in resource allocation patterns. Nutrient availability affected constitutive resistance of poplar to first and fifth instar gypsy moth larvae, which grew much faster on high fertility plants. However, nutrient availability had no effect on constitutive resistance to whitemarked tussock moth. Localized herbivory elicited systemic RIR in poplar within 72 hours. However, the magnitude of RIR was dependent on nutrient availability, with differing effects on the two insect species. Expression of RIR to gypsy moth was most dramatic in the high fertility treatment. In contrast, RIR to whitemarked tussock moth was expressed only in the low fertility treatment. The idiosyncratic nature of effects of nutrient availability on constitutive and induced resistance challenges the value of using insect bioassays as surrogate measures of secondary metabolism for testing allocation models of plant defense, as well as the value of generalized plant defense models for predicting effects of environmental variation on resistance to specific herbivores. These results also suggest that the effects of nutrient availability on the expression of RIR may represent a largely over‐looked source of variation in plant/herbivore interactions.     

Reproductive Biology of an Amphicarpic Annual, Polygonum thunbergii (Polygonaceae): Spatio-temporal Changes in Growth, Structure and Reproductive Components of a Population over an Environmental Gradient

Abstract The process and mechanisms of spatio-temporal changes in growth, population structure, as well as various yield and reproductive components of a population of an amphicarpic annual, Polygonum thunbergii (Polygonaceae), with two reproductive systems (aerial chasmogamous flowers and subterranean cleistogamous flowers) along an environmental gradient (light, moisture and soil nitrogen levels) were investigated in the field.
The results clearly demonstrate that growth and allocation patterns, population structure, and reproductive output of individuals changed sharply along the environmental gradient in response to seasonal and spatial changes in resource availability. Models predicted that light conditions bring about one-sided competition, whereas nutrient conditions in the soil engender two-sided competition. As expected, the degree of one-sided competition was prominent in the case of a planophile, Polygonum thunbergii. Allocation patterns, seed outputs, individual seed size as well as relative energy costs of chasmogamous and cleistogamous seed as affected by light and nutrient levels were also critically analyzed. The most noteworthy finding was that the size of chasmogamous seeds sharply decreased in response to a decrease in the light regime, while cleistogamous seed size remained constant along the gradient. However, relative cost of both chasmogamous and cleistogamous seeds sharply increased with decrease in the light level, reflecting different degrees of environmental stress, biotic interference, or both.     

Fertile forests produce biomass more efficiently

Trees with sufficient nutrition are known to allocate carbon preferentially to aboveground plant parts. Our global study of 49 forests revealed an even more fundamental carbon allocation response to nutrient availability: forests with high-nutrient availability use 58?±?3% (mean?±?SE; 17 forests) of their photosynthates for plant biomass production (BP), while forests with low-nutrient availability only convert 42?±?2% (mean?±?SE; 19 forests) of annual photosynthates to biomass. This nutrient effect largely overshadows previously observed differences in carbon allocation patterns among climate zones, forest types and age classes. If forests with low-nutrient availability use 16?±?4% less of their photosynthates for plant growth, what are these used for? Current knowledge suggests that lower BP per unit photosynthesis in forests with low- versus forests with high-nutrient availability reflects not merely an increase in plant respiration, but likely results from reduced carbon allocation to unaccounted components of net primary production, particularly root symbionts.