A Model of Competition between Trees and Grass for Radiation, Water and Nutrients

Conditions for the coexistence of trees and grass are exploredusing a mathematical model describing plant competition forradiation, water and nutrients. The model describes growth ofboth species in terms of key physiological processes (radiationinterception, photosynthesis, respiration, grazing, litterfall,assimilate partitioning, nutrient uptake and water use) andis used to analyse how species compete by depriving each otherof resources essential for growth. Changes of growth parametersare shown to lead to shifts in species composition (e.g. throughreplacement of one species by another). Competition model, vegetation competition, grass-tree competition, carbon budgets, photosynthesis, assimilate partitioning     

Modelling ozone effects on adult beech trees through simulation of defence, damage, and repair costs: Implementation of the CASIROZ ozone model in the ANAFORE forest model

Ozone affects adult trees significantly, but effects on stem growth are hard to prove and difficult to correlate with the primary sites of ozone damage at the leaf level. To simulate ozone effects in a mechanistic way, at a level relevant to forest stand growth, we developed a simple ozone damage and repair model (CASIROZ model) that can be implemented into mechanistic photosynthesis and growth models. The model needs to be parameterized with cuvette measurements on net photosynthesis and dark respiration. As the CASIROZ ozone sub-model calculates effects of the ozone flux, a reliable representation of stomatal conductance and therefore ozone uptake is necessary to allow implementation of the ozone sub-model. In this case study the ozone sub-model was used in the ANAFORE forest model to simulate gas exchange, growth, and allocation. A preliminary run for adult beech (FAGUS SYLVATICA) under different ozone regimes at the Kranzberg forest site (Germany) was performed. The results indicate that the model is able to represent the measured effects of ozone adequately, and to distinguish between immediate and cumulative ozone effects. The results further help to understand ozone effects by distinguishing defence from damage and repair. Finally, the model can be used to extrapolate from the short-term results of the field study to long-term effects on tree growth. The preliminary simulations for the Kranzberg beech site show that, although ozone effects on yearly growth are variable and therefore insignificant when measured in the field, they could become significant at longer timescales (above 5 years, 5 % reduction in growth). The model offers a possible explanation for the discrepancy between the significant effects on photosynthesis (10 to 30 % reductions simulated), and the minor effects on growth. This appears to be the result of the strong competition and slow growth of the Kranzberg forest, and the importance of stored carbon for the adult beech (by buffering effects on carbon gain). We finally conclude that inclusion of ozone effects into current forest growth and yield models can be an important improvement into their overall performance, especially when simulating younger and less dense forests.     

A Transport-resistance Model of Forest Growth and Partitioning

The transport-resistance approach to dry-matter partitioningis used to construct a model of forest growth The model is atthe stand level for a monoculture of identical trees of thesame age There are five major organ compartments in the modelfoliage, branches, stem, coarse roots, and fine roots and mycorrhizasThe matter in each compartment is further subdivided into menstem,structure, carbon substrate, and nitrogen substrate The modelis driven by daily radiation including day length, ambient CO₂concentration, and daily means of air and soil temperature Thefine roots are provided with constant values of soil mineralnitrogen pools (ammonium and nitrate) from which uptake occursGrowth over about 100 years is simulated for various environmentalconditions and soil mineral nitrogen levels, thinning is alsosimulated Natural tree death occurs within the model Particularattention is paid to dry matter partitioning patterns, and tothe dry matter per stem when death occurs The model is robustand responsive, and provides a framework for further developmentand application to many ecological and environmental scenarios,as well as to some forest management problems Model, forest, growth, partitioning     

Carbon allocation in forest ecosystems

Carbon allocation plays a critical role in forest ecosystem carbon cycling. We reviewed existing literature and compiled annual carbon budgets for forest ecosystems to test a series of hypotheses addressing the patterns, plasticity, and limits of three components of allocation: biomass, the amount of material present; flux, the flow of carbon to a component per unit time; and partitioning, the fraction of gross primary productivity (GPP) used by a component. Can annual carbon flux and partitioning be inferred from biomass? Our survey revealed that biomass was poorly related to carbon flux and to partitioning of photosynthetically derived carbon, and should not be used to infer either. Are component fluxes correlated? Carbon fluxes to foliage, wood, and belowground production and respiration all increased linearly with increasing GPP (a rising tide lifts all boats). Autotrophic respiration was strongly linked to production for foliage, wood and roots, and aboveground net primary productivity and total belowground carbon flux (TBCF) were positively correlated across a broad productivity gradient. How does carbon partitioning respond to variability in resources and environment? Within sites, partitioning to aboveground wood production and TBCF responded to changes in stand age and resource availability, but not to competition (tree density). Increasing resource supply and stand age, with one exception, resulted in increased partitioning to aboveground wood production and decreased partitioning to TBCF. Partitioning to foliage production was much less sensitive to changes in resources and environment. Overall, changes in partitioning within a site in response to resource supply and age were small (<15% of GPP), but much greater than those inferred from global relationships. Across all sites, foliage production plus respiration, and total autotrophic respiration appear to use relatively constant fractions of GPP – partitioning to both was conservative across a broad range of GPP – but values did vary across sites. Partitioning to aboveground wood production and to TBCF were the most variable – conditions that favored high GPP increased partitioning to aboveground wood production and decreased partitioning to TBCF. Do priorities exist for the products of photosynthesis? The available data do not support the concept of priorities for the products of photosynthesis, because increasing GPP increased all fluxes. All facets of carbon allocation are important to understanding carbon cycling in forest ecosystems. Terrestrial ecosystem models require information on partitioning, yet we found few studies that measured all components of the carbon budget to allow estimation of partitioning coefficients. Future studies that measure complete annual carbon budgets contribute the most to understanding carbon allocation.     

Effect of nitrogen deficiency on photosynthesis and the partitioning of 14C-labelled leaf assimilate in unshaded and partially shaded plants of Lolium temulentum

The effect of a deficiency of applied nitrogen on the rate of leaf photosynthesis, and on the subsequent partitioning of ¹⁴C-labelled leaf assimilate between new leaf, stem, tillers and root, was investigated in single plants of Lolium temulentum L., grown normally in controlled environments, or grown with collars shading the base of the plant. The nitrogen deficiency reduced the rate of leaf photosynthesis, increased the retention of assimilate in the leaf, suppressed the export of assimilate to tillers, and generally increased the export of assimilate to roots and to new leaves. Shading the base of the plant generally had little effect on the rate of leaf photosynthesis, reduced the export of assimilate to roots, and increased the export of assimilate to new leaf and to the stem, which elongated when shading was imposed.     

Defoliation,Photosynthetic Rates,and Assimilate Transport in Grapevine Plants

The source-sink relations in grapevine (Vitis vinifera L., var. Rkatsiteli) plants were disturbed by defoliation at different stages of vegetative growth in order to investigate changes in photosynthetic activity and assimilate partitioning. Defoliation was shown to stimulate photosynthesis in the remaining source leaves, enhance the assimilate export, and diminish the midday suppression of photosynthesis. Defoliation created a powerful sink for assimilates, and stimulated their delivery to the affected zone. It is hypothesized that defoliation-induced stress is accompanied by a substantial enhancement of photosynthetic activity and by redistribution of assimilate flows, which enables a sustained supply of assimilates to the sink organs of grapevine plants.__________Translated from Fiziologiya Rastenii, Vol. 52, No. 4, 2005, pp. 507–512.Original Russian Text Copyright © 2005 by Chanishvili, Badridze, Barblishvili, Dolidze.     

The role of sink demand in carbon partitioning and photosynthetic reinvigoration following shoot decapitation

Photosynthesis, growth, and carbon partitioning of vigorous coppice shoots were compared with the slower growing intact shoots of Populus maximowiczii × nigra L. MN9 to determine the relationship between carbon partitioning and photosynthetic rate. Relative height growth rate of coppice shoots was 2.2 times that of intact shoots with net photosynthetic rate 1.9 times that of intact shoots. Coppice leaves exported a larger proportion of newly-fixed assimilate (11% compared with 6%) after a 4-h chase. The greater export from coppice leaves was correlated with a greater proportion of [¹⁴C]-labelled photosynthate deposited as starch in stems 4 cm below the point of label application. Coppice leaf assimilate levels were reduced to 15% that of leaves on intact plants, but coppice leaves had twice the concentration of labelled sucrose. Carbohydrates constituted 55% of the water-soluble [¹⁴C]-labelled photosynthate in leaves of coppice shoots compared with 40% in intact shoots. The results suggest that carbon allocation and partitioning in coppice shoots were altered towards production and export of new assimilate, and support the hypothesis that photosynthetic rate is responsive to sink demand for assimilates.     

Modelling the limits on the response of net carbon exchange to fertilization in a south-eastern pine forest

Using a combination of model simulations and detailed measurements at a hierarchy of scales conducted at a sandhills forest site, the effect of fertilization on net ecosystem exchange (NEE) and its components in 6‐year‐old Pinus taeda stands was quantified. The detailed measurements, collected over a 20‐d period in September and October, included gas exchange and eddy covariance fluxes, sampled for a 10‐d period each at the fertilized stand and at the control stand. Respiration from the forest floor and above‐ground biomass was measured using chambers during the experiment. Fertilization doubled leaf area index (LAI) and increased leaf carboxylation capacity by 20%. However, this increase in total LAI translated into an increase of only 25% in modelled sunlit LAI and in canopy photosynthesis. It is shown that the same climatic and environmental conditions that enhance photosynthesis in the September and October periods also cause an increase in respiration The increases in respiration counterbalanced photosynthesis and resulted in negligible NEE differences between fertilized and control stands. The fact that total biomass of the fertilized stand exceeded 2·5 times that of the control, suggests that the counteracting effects cannot persist throughout the year. In fact, modelled annual carbon balance showed that gross primary productivity (GPP) increased by about 50% and that the largest enhancement in NEE occurred in the spring and autumn, during which cooler temperatures reduced respiration more than photosynthesis. The modelled difference in annual NEE between fertilized and control stands (approximately 200 1;g 2;C 3;m^?2 y^?1) suggest that the effect of fertilization was sufficiently large to transform the stand from a net terrestrial carbon source to a net sink.     

Simulating stand climate, phenology, and photosynthesis of a forest stand with a process-based growth model

In the face of climate change and accompanying risks, forest management in Europe is becoming increasingly important. Model simulations can help to understand the reactions and feedbacks of a changing environment on tree growth. In order to simulate forest growth based on future climate change scenarios, we tested the basic processes underlying the growth model BALANCE, simulating stand climate (air temperature, photosynthetically active radiation (PAR) and precipitation), tree phenology, and photosynthesis. A mixed stand of 53- to 60-year-old Norway spruce (Picea abies) and European beech (Fagus sylvatica) in Southern Germany was used as a reference. The results show that BALANCE is able to realistically simulate air temperature gradients in a forest stand using air temperature measurements above the canopy and PAR regimes at different heights for single trees inside the canopy. Interception as a central variable for water balance of a forest stand was also estimated. Tree phenology, i.e. bud burst and leaf coloring, could be reproduced convincingly. Simulated photosynthesis rates were in accordance with measured values for beech both in the sun and the shade crown. For spruce, however, some discrepancies in the rates were obvious, probably due to changed environmental conditions after bud break. Overall, BALANCE has shown to respond to scenario simulations of a changing environment (e.g., climate change, change of forest stand structure).     

Carbon balance in a monospecific stand of an annual herb <Emphasis Type="Italic">Chenopodium album</Emphasis> at an elevated CO<Subscript>2</Subscript> concentration

Elevated CO₂ enhances carbon uptake of a plant stand, but the magnitude of the increase varies among growth stages. We studied the relative contribution of structural and physiological factors to the CO₂ effect on the carbon balance during stand development. Stands of an annual herb Chenopodium album were established in open-top chambers at ambient and elevated CO₂ concentrations (370 and 700 μmol mol⁻¹). Plant biomass growth, canopy structural traits (leaf area, leaf nitrogen distribution, and light gradient in the canopy), and physiological characteristics (leaf photosynthesis and respiration of organs) were studied through the growing season. CO₂ exchange of the stand was estimated with a canopy photosynthesis model. Rates of light-saturated photosynthesis and dark respiration of leaves as related with nitrogen content per unit leaf area and time-dependent reduction in specific respiration rates of stems and roots were incorporated into the model. Daily canopy carbon balance, calculated as an integration of leaf photosynthesis minus stem and root respiration, well explained biomass growth determined by harvests (r ² = 0.98). The increase of canopy photosynthesis with elevated CO₂ was 80% at an early stage and decreased to 55% at flowering. Sensitivity analyses suggested that an alteration in leaf photosynthetic traits enhanced canopy photosynthesis by 40–60% throughout the experiment period, whereas altered canopy structure contributed to the increase at the early stage only. Thus, both physiological and structural factors are involved in the increase of carbon balance and growth rate of C. album stands at elevated CO₂. However, their contributions were not constant, but changed with stand development.     

陆地碳循环研究中植物生理生态过程模拟进展

植物生理生态过程的模拟是陆地碳循环模型研究中的关键过程之一,就与碳循环过程密切相关的3个关键的植物生理生态过程;光合作用,碳分配和物候等过程的数学模式进行分类：（1）光合作用模式,根据光合作用模式基础的不同把光合作用模式分为：半经验模式;机理模式和使用卫星遥感资料的模式等;（2）植物碳分配模式介绍了功能平衡模式;运输-阻力模式;光合作用与生长模式;环境反应模式和大尺度生态系统模式等5类。（3）植被物候模式;根据观测和参数化方法的不同可以将现有的物候数值模式分为两类;观测统计模式和使用卫星遥感资料的物候模式,对各类模式的主要控制方程,研究进展和应用分别进行了简要评述。     

Plant respiration: Controlled by photosynthesis or biomass?

Two simplifying hypotheses have been proposed for whole‐plant respiration. One links respiration to photosynthesis; the other to biomass. Using a first‐principles carbon balance model with a prescribed live woody biomass turnover, applied at a forest research site where multidecadal measurements are available for comparison, we show that if turnover is fast the accumulation of respiring biomass is low and respiration depends primarily on photosynthesis; while if turnover is slow the accumulation of respiring biomass is high and respiration depends primarily on biomass. But the first scenario is inconsistent with evidence for substantial carry‐over of fixed carbon between years, while the second implies far too great an increase in respiration during stand development—leading to depleted carbohydrate reserves and an unrealistically high mortality risk. These two mutually incompatible hypotheses are thus both incorrect. Respiration is not linearly related either to photosynthesis or to biomass, but it is more strongly controlled by recent photosynthates (and reserve availability) than by total biomass.     

Carbon partitioning in Arabidopsis thaliana is a dynamic process controlled by the plants metabolic status and its circadian clock

Plant growth involves the coordinated distribution of carbon resources both towards structural components and towards storage compounds that assure a steady carbon supply over the complete diurnal cycle. We used ¹⁴CO₂ labelling to track assimilated carbon in both source and sink tissues. Source tissues exhibit large variations in carbon allocation throughout the light period. The most prominent change was detected in partitioning towards starch, being low in the morning and more than double later in the day. Export into sink tissues showed reciprocal changes. Fewer and smaller changes in carbon allocation occurred in sink tissues where, in most respects, carbon was partitioned similarly, whether the sink leaf assimilated it through photosynthesis or imported it from source leaves. Mutants deficient in the production or remobilization of leaf starch exhibited major alterations in carbon allocation. Low‐starch mutants that suffer from carbon starvation at night allocated much more carbon into neutral sugars and had higher rates of export than the wild type, partly because of the reduced allocation into starch, but also because of reduced allocation into structural components. Moreover, mutants deficient in the plant's circadian system showed considerable changes in their carbon partitioning pattern suggesting control by the circadian clock.     

Lack of fructose 2,6‐bisphosphate compromises photosynthesis and growth in Arabidopsis in fluctuating environments

The balance between carbon assimilation, storage and utilisation during photosynthesis is dependent on partitioning of photoassimilate between starch and sucrose, and varies in response to changes in the environment. However, the extent to which the capacity to modulate carbon partitioning rapidly through short‐term allosteric regulation may contribute to plant performance is unknown. Here we examine the physiological role of fructose 2,6‐bisphosphate (Fru‐2,6‐P₂) during photosynthesis, growth and reproduction in Arabidopsis thaliana (L.). In leaves this signal metabolite contributes to coordination of carbon assimilation and partitioning during photosynthesis by allosterically modulating the activity of cytosolic fructose‐1,6‐bisphosphatase. Three independent T‐DNA insertional mutant lines deficient in 6‐phosphofructo‐2‐kinase/fructose‐2,6‐bisphosphatase (F2KP), the bifunctional enzyme responsible for both the synthesis and degradation of Fru‐2,6‐P₂, lack Fru‐2,6‐P₂. These plants have normal steady‐state rates of photosynthesis, but exhibit increased partitioning of photoassimilate into sucrose and have delayed photosynthetic induction kinetics. The F2KP‐deficient plants grow normally in constant environments, but show reduced growth and seed yields relative to wildtype plants in fluctuating light and/or temperature. We conclude that Fru‐2,6‐P₂ is required for optimum regulation of photosynthetic carbon metabolism under variable growth conditions. These analyses suggest that the capacity of Fru‐2,6‐P₂ to modulate partitioning of photoassimilate is an important determinant of growth and fitness in natural environments.     

Linking drought legacy effects across scales: From leaves to tree rings to ecosystems

Severe drought can cause lagged effects on tree physiology that negatively impact forest functioning for years. These “drought legacy effects” have been widely documented in tree‐ring records and could have important implications for our understanding of broader scale forest carbon cycling. However, legacy effects in tree‐ring increments may be decoupled from ecosystem fluxes due to (a) postdrought alterations in carbon allocation patterns; (b) temporal asynchrony between radial growth and carbon uptake; and (c) dendrochronological sampling biases. In order to link legacy effects from tree rings to whole forests, we leveraged a rich dataset from a Midwestern US forest that was severely impacted by a drought in 2012. At this site, we compiled tree‐ring records, leaf‐level gas exchange, eddy flux measurements, dendrometer band data, and satellite remote sensing estimates of greenness and leaf area before, during, and after the 2012 drought. After accounting for the relative abundance of tree species in the stand, we estimate that legacy effects led to ~10% reductions in tree‐ring width increments in the year following the severe drought. Despite this stand‐scale reduction in radial growth, we found that leaf‐level photosynthesis, gross primary productivity (GPP), and vegetation greenness were not suppressed in the year following the 2012 drought. Neither temporal asynchrony between radial growth and carbon uptake nor sampling biases could explain our observations of legacy effects in tree rings but not in GPP. Instead, elevated leaf‐level photosynthesis co‐occurred with reduced leaf area in early 2013, indicating that resources may have been allocated away from radial growth in conjunction with postdrought upregulation of photosynthesis and repair of canopy damage. Collectively, our results indicate that tree‐ring legacy effects were not observed in other canopy processes, and that postdrought canopy allocation could be an important mechanism that decouples tree‐ring signals from GPP.     

Carbon metabolism in spinach leaves as affected by leaf age and phosphorus and sulfur nutrition

Spinach (Spinacea oleracea) plants were grown either continuously on complete nutrient solutions or for 2 weeks on media deficient in phosphate or sulfate. To characterize leaf carbohydrate metabolism, levels of phosphorylated intermediates, activities of enzymes involved in photosynthetic carbon metabolism, contents of soluble and acid hydrolyzable sugars were measured in leaves differing in age and mineral status and related to leaf rates of photosynthesis and assimilate partitioning. Concentrations of metabolites—particularly those which are preferentially compartmented in the cytosol—decreased from young to old leaves and were lowest in old phosphate starved leaves. Nutrient deficiency showed comparable effects on stromal and cytosolic intermediates. Whole leaf ATP to ADP ratios were dependent on the growth regime, but did not much change with leaf age. The assimilatory force increased in all leaves suffering from mineral deficiency; the assimilatory force was low when photosynthesis was high and vice versa. Sugars accumulated although enzyme activities were decreased under deficiency. The results show that growth of P- and S-starved plants is not limited by photosynthetic reactions.     

Age-dependent response of boreal forest to temperature and rainfall variability

We used measurements of CO₂ exchange by six Canadian boreal forest stands to determine how sites of contrasting age respond to interannual temperature and precipitation variation. The stands ranged in age (time since last burn) from 4 to 155 years. The study included years that were anomalously cool and moist, warm and dry, cool and dry, and warm and moist. Warmer than average springs accelerated the onset of older stand (> 20 years) photosynthesis by as much as 37 days and younger stand (< 20 years) photosynthesis by as much as 25 days. The warm–dry year resulted in a marked reduction of mid‐summer CO₂ uptake by the younger, but not older, stands. The mid‐summer decline in young stand photosynthesis reflected the combination of warmth and drought; similar declines were not observed during the cool–dry or warm–moist years. The annual carbon gain by the oldest stands was greatest during the warm–dry year as a result of the expanded growing season. The annual carbon gain by the youngest stands was consistent from year to year, largely as a result of offsets between increased spring photosynthesis and reduced mid‐summer photosynthesis during the warm–dry year. Night‐time CO₂ efflux increased by 2–29% during the warm–moist year relative to the warm–dry year. This increase was not systematically related to age. We conclude that the major source of interannual CO₂ exchange variation at the landscape scale is the ability of older, evergreen canopies to respond to warm springs by advancing the onset of photosynthesis. Drought‐related reductions in photosynthesis, moisture‐driven respiratory losses, and the effects of spring warmth on young‐stand photosynthesis are of secondary importance. The advantage conferred on older, evergreen stands during warmer years carries implications for the possible effects of climate change on boreal forest succession. Warmer temperatures may increase the competitive advantage of late successional species relative to early successional species, and also the incidence of fire, thereby accelerating plant succession and shortening the fire‐return interval.     

The proton-sucrose symport

The heterotrophic tissues of the plant are dependent upon carbon and nitrogen import for normal growth and development. In general, oxidized forms of these essential elements are reductively assimilated in the leaf and, subsequently, sucrose and amino acids are transported to the heterotrophic cells in a process known as assimilate partitioning. In many plant species, a critical component of the assimilate partitioning pathway is the proton-sucrose symport. This active transport system couples sucrose translocation across the plasma membrane to the proton motive force generated by the H⁺-pumping ATPase. To date, the proton-sucrose symport is the only known system that can account for sucrose accumulation in the vascular tissue of the plant. This review focuses on recent advances describing the transport properties and bioenergetics of the proton-sucrose symport.     

森林碳库特征及驱动因子分析研究进展

森林碳库作为全球碳库的重要组成部分,在区域以及全球碳循环中发挥重要作用。森林生态系统有机碳库主要由3部分组成:活植物碳库、土壤有机质碳库和死植物体碳库。各碳库时空差异很大,使研究森林碳储存机制存在很大的不确定性。在全球或者区域尺度上,森林生物量(与森林碳储量密切相关)呈现出清晰的分布格局,但对于这些格局的驱动因子与地上生物量的相关关系和尺度外推程度尚有很大的争议。分别讨论了气候、土壤、地形和生物因子对地上生物量的影响;而后,从各生态因子与地上生物量之间的直接或者间接关系入手,分析了各因子与生物量之间的交互作用,认为目前运用相同的处理方式来研究环境因子和生物因子对森林生物量的影响是不合适的。文章最后介绍两种新的研究方法:层次模型和生境分类,以期用来重新评估生态因子对森林生物量的影响,有助于更准确的了解森林碳储存机制。     

Photosynthesis and assimilate partitioning between carbohydrates and isoprenoid products in vegetatively active and dormant guayule: physiological and environmental constraints on rubber accumulation in a semiarid shrub

The stems and roots of the semiarid shrub guayule, Parthenium argentatum, contain a significant amount of natural rubber. Rubber accumulates in guayule when plants are vegetatively and reproductively dormant, complicating the relationship between growth/reproduction and product synthesis. To evaluate the factors regulating the partitioning of carbon to rubber, carbon assimilation and partitioning were measured in guayule plants that were grown under simulated summer‐ and winter‐like conditions and under winter‐like conditions with CO₂ enrichment. These conditions were used to induce vegetatively active and dormant states and to increase the source strength of vegetatively dormant plants, respectively. Rates of CO₂ assimilation, measured under growth temperatures and CO₂, were similar for plants grown under summer‐ and winter‐like conditions, but were higher with elevated CO₂. After 5 months, plants grown under summer‐like conditions had the greatest aboveground biomass, but the lowest levels of non‐structural carbohydrates and rubber. In contrast, the amount of resin in the stems was similar under all growth conditions. Emission of biogenic volatile compounds was more than three‐fold higher in plants grown under summer‐ compared with winter‐like conditions. Taken together, the results show that guayule plants maintain a high rate of photosynthesis and accumulate non‐structural carbohydrates and rubber in the vegetatively dormant state, but emit volatile compounds at a lower rate when compared with more vegetatively active plants. Enrichment with CO₂ in the vegetatively dormant state increased carbohydrate content but not the amount of rubber, suggesting that partitioning of assimilate to rubber is limited by sink strength in guayule.