Carbon and nitrogen assimilation in relation to yield: mechanisms are the key to understanding production systems

Improved understanding of crop production systems in relation to N-supply has come from a knowledge of basic plant biochemistry and physiology. Gene expression leads to protein synthesis and the formation of metabolic systems; the ensuing metabolism determines the capacity for growth, development and yield production. This constitutes the genetic potential. These processes set the requirements for the supply of resources. The interactions between carbon dioxide (CO(2)) and nitrate () assimilation and their dynamics are of key importance for crop production. In particular, an adequate supply of, its assimilation to amino acids (for which photosynthesized carbon compounds are required) and their availability for protein synthesis, are essential for metabolism. An adequate supply of stimulates leaf growth and photosynthesis, the former via cell growth and division, the latter by larger contents of components of the light reactions, and those of CO(2) assimilation and related processes. If the supply of resources exceeds the demand set by the genetic potential then production is maximal, but if it is less then potential is not reached; matching resources to potential is the aim of agriculture. However, the connection between metabolism and yield is poorly quantified. Biochemical characteristics and simulation models must be better used and combined to improve fertilizer-N application, efficiency of N-use, and yields. Increasing N-uptake at inadequate N-supply by increasing rooting volume and density is feasible, increasing affinity is less so. It would increase biomass and N/C ratio. With adequate N, at full genetic potential, more C-assimilation per unit N would increase biomass, but energy would be limiting at full canopy. Increasing C-assimilation per unit N would increase biomass but decrease N/C at both large and small N-supply. Increasing production of all biochemical components would increase biomass and demand for N, and maintain N/C ratio. Changing C- or N-assimilation requires modifications to many processes to effect improvements in the whole system; genetic engineering/molecular biological alterations to single steps in the central metabolism are unlikely to achieve this, because targets are unclear, and also because of the complex interactions between processes and environment. Achievement of the long-term objectives of improving crop N-use and yield with fewer inputs and less pollution, by agronomy, breeding or genetic engineering, requires a better understanding of the whole system, from genes via metabolism to yield.     

A semimechanistic model predicting the growth and production of the bioenergy crop Miscanthus×giganteus: description, parameterization and validation

Biomass based bioenergy is promoted as a major sustainable energy source which can simultaneously decrease net greenhouse gas emissions. Miscanthus × giganteus ( M. × giganteus ), a C₄ perennial grass with high nitrogen, water, and light use efficiencies, is regarded as a promising energy crop for biomass production. Mathematical models which can accurately predict M. × giganteus biomass production potential under different conditions are critical to evaluate the feasibility of its production in different environments. Although previous models based on light-conversion efficiency have been shown to provide good predictions of yield, they cannot easily be used in assessing the value of physiological trait improvement or ecosystem processes. Here, we described in detail the physical and physiological processes of a previously published generic mechanistic eco-physiological model, WIMOVAC, adapted and parameterized for M. × giganteus . Parameterized for one location in England, the model was able to realistically predict daily field diurnal photosynthesis and seasonal biomass at a range of other sites from European studies. The model provides a framework that will allow incorporation of further mechanistic information as it is developed for this new crop.     

Seasonal pattern of abundance of honeyeaters and their resources in heathland areas near Sydney

Abundances of honeyeaters, flowers and flying insects, the daily nectar-energy production per flower and the average size of flying insects were estimated every three weeks for circles of radius 20 m located in three heathland areas near Sydney. Seasonal fluctuations in honeyeater density showed no apparent relationship with seasonal fluctuations in nectar-energy productivity or in biomass of flying insects. Variation between circles in honeyeater density was also unrelated to spatial variation in energy productivity and insect biomass. The relatively low incidence of nectar-feeding and high incidence of flying exhibited by birds observed during troughs in nectar-energy production suggest that many of these birds are transient and that their density may consequently be unrelated to local conditions.     

Role of ageing and temperature in shaping reaction norms and fecundity functions in insects

The existing energy partitioning models assume that fecundity is constant throughout adult life. In insects, however, fecundity is a triangular function of time: after maturation, it initially sharply increases and after reaching its maximum it slowly declines as the mother ages. These models also fail to explain that empirical data generally indicate an increase in juvenile growth rate caused by improvement in food quality results in larger adults, whereas that caused by an increase in ambient temperature results in smaller adults. This ‘life history puzzle’ has worried many biologists for a long time. An energy‐partitioning model for insects is presented with soma and gonads as its components, which – contrary to other models – assumes ageing of soma. This model explains the triangular shape of the fecundity function, and also offers an explanation of the ‘life history puzzle’. The differential response in adult size to changes in food quality and temperature in nature may result from the differential responses of our model’s parameters to changes in these environmental parameters. Better food quality results in bigger adults, because food quality affects the assimilation rate, but not the rate of conversion of gonadal biomass into offspring, or the rate of senescence. In contrast, an increase in temperature speeds up all the processes. That is, temperature affects the assimilation rate, the conversion rate of gonadal biomass into offspring, and the rate of senescence equally. Therefore, an increase in temperature results in larger or smaller adults, depending on the shape of the senescence function.     

Atmospheric CO2 enrichment alters energy assimilation, investment and allocation in Xanthium strumarium

Energy-use efficiency and energy assimilation, investment and allocation patterns are likely to influence plant growth responses to increasing atmospheric CO2 concentration ([CO2]). Here, we describe the influence of elevated [CO2] on energetic properties as a mechanism of growth responses in Xanthium strumarium. Individuals of X. strumarium were grown at ambient or elevated [CO2] and harvested. Total biomass and energetic construction costs (CC) of leaves, stems, roots and fruits and percentage of total biomass and energy allocated to these components were determined. Photosynthetic energy-use efficiency (PEUE) was calculated as the ratio of total energy gained via photosynthetic activity (Atotal) to leaf CC. Elevated [CO2] increased leaf Atotal, but decreased CC per unit mass of leaves and roots. Consequently, X. strumarium individuals produced more leaf and root biomass at elevated [CO2] without increasing total energy investment in these structures (CCtotal). Whole-plant biomass was associated positively with PEUE. Whole-plant construction required 16.1% less energy than modeled whole-plant energy investment had CC not responded to increased [CO2]. As a physiological mechanism affecting growth, altered energetic properties could positively influence productivity of X. strumarium, and potentially other species, at elevated [CO2].     

Direct and size-mediated effects of temperature and ration-dependent growth rates on energy reserves in juvenile anadromous alewives (Alosa pseudoharengus)

Growth rate and energy reserves are important determinants of fitness and are governed by endogenous and exogenous factors. Thus, examining the influence of individual and multiple stressors on growth and energy reserves can help estimate population health under current and future conditions. In young anadromous fishes, freshwater habitat quality determines physiological state and fitness of juveniles emigrating to marine habitats. In this study, the authors tested how temperature and food availability affect survival, growth and energy reserves in juvenile anadromous alewives (Alosa pseudoharengus), a forage fish distributed along the eastern North American continent. Field-collected juvenile anadromous A. pseudoharengus were exposed for 21 days to one of two temperatures (21°C and 25°C) and one of two levels of food rations (1% or 2% tank biomass daily) and compared for differences in final size, fat mass-at-length, lean mass-at-length and energy density. Increased temperature and reduced ration both led to lower growth rates, and the effect of reduced ration was greater at higher temperature. Fat mass-at-length decreased with dry mass, and energy density increased with total length, suggesting size-based endogenous influences on energy reserves. Lower ration also directly decreased fat mass-at-length, lean mass-at-length and energy density. Given the fitness implications of size and energy reserves, temperature and food availability should be considered important indicators of nursery habitat quality and incorporated in A. pseudoharengus life-history models to improve forecasting of population health under climate change.     

The effects of control on the biomass, carbohydrate content and bud reserves of bracken (Pteridium aquilinum (L.) Kuhn), and an evaluation of a bracken growth model

This paper examines the initial effects of bracken control on frond numbers and biomass, and the biomass, carbohydrate reserves and bud densities of bracken stands cut once per year, twice per year, subject to a single application of asulam or left untreated. The seasonal dynamics of these parameters are displayed; carbohydrate and biomass are both removed from the rhizome system to produce frond tissue, and are replenished at the end of the growing season. Asulam application reduced densities of both active and dormant buds, and both frond biomass and density. It did not significantly reduce rhizome biomass or carbohydrate reserves in the two years after treatment. Cutting, either once or twice per year reduced both rhizome biomass and rhizome carbohydrate reserves, as well as bud densities, though the latter were reduced in proportion to biomass. Cutting twice a year reduced the production of fronds, both in numbers and biomass. The collected data were used to evaluate a model of bracken growth, and subsequently to improve estimates of some of the model parameters. The model simulations of control treatments were compared to field data. The effects of cutting once per year and spraying with asulam were predicted accurately, but the bracken stand was more resilient to cutting twice per year than would be expected from model predictions. The combination of cutting and spraying is discussed as a potential tool in land management and the deficiencies of the model are discussed in relation to the need for future research into the biology of bracken.     

A Dynamic Energy Budget model based on partitioning of net production

We formulate a Dynamic Energy Budget (DEB) model for the growth and reproduction of individual organisms based on partitioning of net production (i.e. energy acquisition rate minus maintenance rate) between growth and energy reserves. Reproduction uses energy from reserves. The model describes both feeding and non-feeding stages, and hence is applicable to embryos (which neither feed nor reproduce), juveniles (which feed but do not reproduce), and adults (which commonly both feed and reproduce). Embryonic growth can have two forms depending on the assumptions for acquisition of energy from yolk. By default, when the energy acquisition rate exceeds the maintenance rate, a fixed proportion of the resulting net production is spent on growth (increase in structural biomass), and the remaining portion is channelled to the reserves. Feeding organisms, however, modulate their allocation of net production energy in response to their total energy content (energy in the reserves plus energy bounded to structural biomass). In variable food environment an organism alternates between periods of growth, no-growth, and balanced-growth. In the latter case the organism adopts an allocation strategy that keeps its total energy constant. Under constant environmental conditions, the growth of a juvenile is always of von Bertalanffy type. Depending on the values of model parameters there are two long-time possibilities for adults: (a) von Bertalanffy growth accompanied by reproduction at a rate that approaches zero as the organism approaches asymptotic size, or (b) abrupt cessation of growth at some finite time, following which, the rate of reproduction is constant. We illustrate the model's applicability in life history theory by studying the optimum values of the energy allocation parameters for constant environment and for each of the dynamic regimes described above. Received: 11 May 1998 / Revised version: 18 February 2000 / Published online: 4 October 2000     

Ecological and mathematical considerations on self-thinning in even-aged pure stands

It is emphasized in growth analysis of self-thinning populations that relative mortality rate pertains to the difference between relative growth rates and net assimilation rates, each of which are definable on a mean plant size basis or on a biomass basis. The time trends of the ratio of relative mortality rate to relative growth rates to be expected according to Tadaki's, Shinozaki's and Hozumi's models are compared with that of the eastern white pine population, and a good agreement is exhibited. As an alternative to Hozumi's model, a new model is constructed to unite the logistic theory of plant growth and the 3/2 power law concerning self-thinning, which so far have usually been applied independently to growth analysis. To construct the model the following assumptions are made: the fundamental equation to relate mean plant weight with density in self-thinning population proposed by Shinozaki, and a special population with a specific initial density which follows thew-p trajectory of the 3/2 power law type and has an exponential decrease in its density with biological time. Properties of the model are examined from ecological and mathematical viewpoints.     

Bi-trophic food chain dynamics with multiple component populations

Food web models describe the patterns of material and energy flow in communities. In classical food web models the state of each population is described by a single variable which represents, for instance, the biomass or the number of individuals that make up the population. However, in a number of models proposed recently in the literature the individual organisms consist of two components. In addition to the structural component there is an internal pool of nutrients, lipids or reserves. Consequently the population model for each trophic level is described by two state variables instead of one. As a result the classical predator-prey interaction formalisms have to be revised. In our model time budgets with actions as searching and handling provide the formulation of the functional response for both components. In the model, assimilation of the ingested two prey components is done in parallel and the extracted energy is added to a predators reserve pool. The reserves are used for vital processes; growth, reproduction and maintenance. We will explore the top-down modelling approach where the perspective is from the community. We will demonstrate that this approach facilitates a check on the balance equations for mass and energy at this level of organization. Here it will be shown that, if the individual is allowed to shrink when the energy reserves are in short to pay the maintenance costs, the growth process has to be 100% effective. This is unrealistic and some alternative model formulations are discussed. The long-term dynamics of a microbial food chain in the chemostat are studied using bifurcation analysis. The dilution rate and the concentration of nutrients in the reservoir are the bifurcation parameters. The studied microbial bi-trophic food chain with two-component populations shows chaotic behaviour.     

Spatially implicit approaches to understand the manipulation of mating success for insect invasion management

Recent work indicates that Allee effects (the positive relationship between population size and per capita growth rate) are critical in determining the successful establishment of invading species. Allee effects may create population thresholds, and failure to establish is likely if invading populations fall below these thresholds. There are many mechanisms that may contribute to Allee effects, but mate-location failure is a common cause in sexually reproducing insects. Consequently, mate-location failure represents a type of “weak link” that may be enhanced in order to achieve eradication of insect populations during the early stages of invasion. In this paper, spatially implicit models that account for mating behavior of both sexes are used to explore the enhancement of mate-location failure in pest eradication programs. Distinct from the previous studies, the Allee effect emerges from a mechanistic representation of mate-location failure in our model. Three specific eradication strategies, sterile insect release, mass-trapping, and mating disruption, are incorporated into the model and tested for their ability to depress population growth during the early stages of invasions. We conducted simulations with the model parameterized to represent two types of insects: Coleopteran-like insects which are long-lived and capable of multiple matings, but have low daily reproductive rates, and Lepidopteran-like insects which are capable of mating only once per generation, have an ephemeral reproductive stage, and have high reproductive rates. Simulations indicated that: (1) many insect pests are more likely to be eradicated than had been previously predicted by classic models which do not account for mate-finding difficulties, (2) for Lepidopteran-like insects, mass-trapping has the greatest potential for eradication among the three methods when a large number of traps can be installed, although mating-disruption will be the most effective if we can anticipate confusion or trail-masking mechanisms of disruption, and (3) populations of Coleopteran-like insects may be most effectively eradicated using the sterile male release method. Though more detailed models should be tailored for individual species, we expect that the spatially implicit approaches outlined in this paper can be widely adapted to study the efficiency of various eradication approaches in sparse conditions. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

WBE 模型及其在生态学中的应用：研究概述

介绍了WBE模型,综述了该模型在生态学中的应用进展。WBE模型,以及以该模型为基础的MTE模型,假设生物体为自相似分形网络结构,提出代谢速率和个体大小之间存在3/4指数关系,分别预测了从个体到生物圈多个尺度上的生物属性之间的异速生长关系,而且部分得到了验证。WBE模型的应用涵盖了个体组织生物量、年生长率,种群密度和生态系统单位面积产量、能量流动率等多个方面;即使在生物圈大尺度上,WBE模型也可用来预测试验中无法直接测量的特征变量的属性,如全球碳储量的估算等。至今,关于WBE和MTE模型仍然存在各种褒贬争论,讨论焦点主要集中于模型建立的前提假设以及权度指数的预测。今后的研究工作应规范试验技术和方法,考虑物种多样性和环境等因素的影响,提出符合各类生物的模型结构体系,使其具有更广泛的应用性和预测性。     

Evaluating soil microbial carbon use efficiency explicitly as a function of cellular processes: implications for measurements and models

Carbon use efficiency (CUE), the proportion of carbon (C) consumed by microbes that is converted into biomass, is an important parameter for soil C models with explicit microbial controls. While often considered as a single parameter, CUE is an emergent property of multiple microbial processes, including assimilation efficiency, biomass-specific respiration, enzyme production, and respiratory costs of enzyme production. These processes occur over variable time scales and imply different fates for C, and the same emergent CUE value can result when C is allocated in fundamentally different ways (e.g. a high investment in enzyme production vs. a high assimilation cost). We developed a model that represents the individual processes underlying emergent CUE to test how shifts in microbial allocation alter equilibrium soil C pool sizes. We found that an increase in emergent CUE that results from a change in assimilation efficiency, biomass specific respiration, or respiration costs from enzyme production causes soil organic C (SOC) to decline, while the same change in emergent CUE resulting from a change in enzyme production causes SOC to increase. We also used the model to test the sensitivity of CUE from isotopic C tracer estimates to changes in microbial allocation processes. We found that these estimates do not account for the same microbial processes represented by emergent CUE in models. We propose that considering microbial processes explicitly rather than representing CUE as a single parameter can improve data-model integration. In addition, modeling microbial processes explicitly will account for a wider range of possible outcomes from shifts in microbial C allocation, such as when increased SOC results from increasing CUE.     

Energy flow in four different field populations of Ancylus fluviatilis (Gastropoda-Basommatophora)

Summary Four brook-dwelling populations of Ancylus fluviatilis were studied. Data on life-cycle, specimen number, biomass and egg capsule biomass are reported. By means of experimental data on ingestion, assimilation, growth and egg-production, energy balances are calculated for whole populations.Mean annual biomass turned out to be 26.58–64.88 mg C/m². Mean annual biomass turn-over ratio (production/biomass) was 2.64–5.95. Cohort biomass turnover ratio was 4.11–6.09 for the whole population. Young specimens show considerably higher production efficiencies than adult ones. Production by the smallest size classes is a very considerable part of whole population production. Net production efficiency of the whole population was 11.2–11.8%, gross production efficiency 5.6–6.5%. An important factor for variation between different populations is temperature, but other factors, such as food supply, may be important, too. Experimental production estimates were compaerd with the Allen curve method and the instantaneous growth method.Differences due to different methods of production measurement are discussed. The range of values for different efficiencies lies within the range reported for other freshwater snails. Interpopulation variation in turnover ratios nearly equals variation reported for different species of univoltine snails.Supported by the Deutsche Forschungsgemeinschaft.     

Studies on the chemical ecology of Biomphalaria glabrata: The effects of chemical conditioning by the snails kept at various densities on their growth and metabolism

It has been shown that heterotypically conditioned media (Het. C.M.) produced by lettuce fed snails maintained at various densities can influence the allometric growth and the rates of specific growth, ingestion, egestion, assimilation and heart beat of isolated assay snails. The growth rate of the assay snails was enhanced by increasing the density of snails producing the Het. C.M. to an optimum threshold of one snail per 100 ml. Further increases in density were followed by a decline in their growth rate. The higher growth achieved by the assay snails in the optimum treatment can be attributed to the fact that their specific growth rates over the first three weeks of the experiment were higher than those of snails in other treatments.
Ingestion and assimilation correlated well with specific growth rates. Percentage assimilation values are high, ranging from 78.4 to 89.6%. The onset of reproductive activity is followed by a dramatic decline from approximately 26 % to 5 % in the gross efficiency of utilization of food for somatic growth.
Snails not receiving Het. C.M. from the outer tanks have a relatively higher ratio of specific growth rate in weight to length than was the case with snails in the other treatments. The heart beat rates of snails in the Het. C.M. produced by snails at densities of 80 to 160 snails per 41 are significantly lower than those of snails at other densities. The possible causation and adaptive significance of these effects are discussed.     

Gluttony in migratory waders – unprecedented energy assimilation rates in vertebrates

Maximum energy assimilation rate has been implicated as a constraint on maximal sustained energy expenditure, on biomass production, and in various behavioural and life history models. Data on the upper limit to energy assimilation rate are scarce, and the factors that set the limit remain poorly known. We studied migratory waders in captivity, given unlimited food supply around the clock. Many of these waders assimilated energy at rates of seven to ten times basal metabolism, exceeding maximum rates reported for vertebrates during periods of high energy demand, for example during reproduction and in extreme cold. One factor allowing the high energy assimilation rates may be that much of the assimilated energy is stored and not concomitantly expended by muscles or other organs. The remarkable digestive capacity in waders is probably an adaptation to long and rapid migrations, putting a premium on high energy deposition rates. The upper limit to daily energy assimilation in vertebrates is clearly higher than hitherto believed, and food availability, total daily feeding time and, possibly, the fate of assimilated energy may be important factors to take into account when estimating limits to energy budgets in animals.     

Energy density and its variation in space limit species richness of boreal forest birds

Aim An area’s ability to support species may be dependent not only on the total amount of available energy it contains but also on energy density (i.e. available energy per unit area). Acknowledging these two aspects of energy availability may increase mechanistic understanding of how increased energy availability results in increased species richness. We studied the relationship between energy density, its variation in space and boreal forest bird species richness and investigated two possible mechanisms: (1) metabolic constraints of organisms, and (2) increased resource availability for specialists. Location Protected areas in Finland’s boreal forest. Methods We tested whether bird species richness was best determined by total energy availability in an area or by energy density and its variation within the area, before and after including bird abundance in the models. We evaluated two main explanatory variables: tree growth reflecting the rate of energy production and tree volume as a measure of biomass. In addition, we modelled individual species’ responses to energy density and its variation, and evaluated the prediction of the metabolic constraints hypothesis that small species are limited by energy density whereas large species are limited by total energy availability in the area. Results Energy density and its variation were good predictors of species richness: together with abundance they explained 84% of variation in species richness (compared with 74% for abundance alone). Pure metabolic constraints were unlikely to explain this relationship. Instead, the mechanism probably involved increased habitat heterogeneity benefiting specialist species. Total energy availability was also an important factor determining species richness but its effect was indirect via abundance. Main conclusions Our results corroborate the importance of energy availability as a driver of species richness in forest bird communities, and they indicate that energy density and its variation in the landscape strongly influence species richness even after accounting for abundance.     

Enrichment in a general class of stoichiometric producer-consumer population growth models

This paper presents the derivation and partial analysis of a general producer-consumer model. The model is stoichiometric in that it includes the growth constraints imposed by species-specific biomass carbon to nutrient ratios. The model unifies the approaches of other studies in recent years, and is calibrated from an extensive review of the algae-Daphnia literature. Numerical simulations and bifurcation analysis are used to examine the impact of energy enrichment under nutrient and stoichiometric constraints. Our results suggest that the variety of system responses previously cited for related models can be attributed to the size of the total system nutrient pool, which is here assumed fixed. New, more complicated bifurcation sequences, such as multiple homoclinic bifurcations, are demonstrated as well. The mechanistic basis of the model permits us to show the robustness of the system’s dynamics subject to alternate approaches to modeling producer and consumer biomass production.