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1.  Genetic differences in fruit-set patterns are determined by differences in fruit sink strength and a source : sink threshold for fruit set 
Annals of Botany  2009;104(5):957-964.
Background and Aims
Fruit set in indeterminate plant species largely depends on the balance between source and sink strength. Plants of these species show fluctuations in fruit set during the growing season. It was tested whether differences in fruit sink strength among the cultivars explained the differences in fruit-set patterns.
Capsicum was chosen as a model plant. Six cultivars with differences in fruit set, fruit size and plant growth were evaluated in a greenhouse experiment. Fruit-set patterns, generative and vegetative sink strength, source strength and the source : sink ratio at fruit set were determined. Sink strength was quantified as potential growth rate. Fruit set was related to total fruit sink strength and the source : sink ratio. The effect of differences observed in above-mentioned parameters on fruit-set patterns was examined using a simple simulation model.
Key Results
Sink strengths of individual fruits differed greatly among cultivars. Week-to-week fruit set in large-fruited cultivars fluctuated due to large fluctuations in total fruit sink strength, but in small-fruited cultivars, total fruit sink strength and fruit set were relatively constant. Large variations in week-to-week fruit set were correlated with a low fruit-set percentage. The source : sink threshold for fruit set was higher in large-fruited cultivars. Simulations showed that within the range of parameter values found in the experiment, fruit sink strength and source : sink threshold for fruit set had the largest impact on fruit set: an increase in these parameters decreased the average percentage fruit set and increased variation in weekly fruit set. Both were needed to explain the fruit-set patterns observed. The differences observed in the other parameters (e.g. source strength) had a lower effect on fruit set.
Both individual fruit sink strength and the source : sink threshold for fruit set were needed to explain the differences observed between fruit-set patterns of the six cultivars.
PMCID: PMC2749527  PMID: 19643909
Fruit-set patterns; fruit sink strength; source : sink ratio; threshold for fruit set; Capsicum annuum; cultivars
2.  Correlation between dynamic tomato fruit-set and source–sink ratio: a common relationship for different plant densities and seasons? 
Annals of Botany  2010;107(5):805-815.
Background and Aims
It is widely accepted that fruit-set in plants is related to source–sink ratio. Despite its critical importance to yield, prediction of fruit-set remains an ongoing problem in crop models. Functional–structural plant models are potentially able to simulate organ-level plasticity of plants. To predict fruit-set, the quantitative link between source–sink ratio and fruit-set probability is analysed here via a functional–structural plant model, GreenLab.
Two experiments, each with four plant densities, were carried out in a solar greenhouse during two growth seasons (started in spring and autumn). Dynamic fruit-set probability was estimated by frequent observation on inflorescences. Source and sink parameter values were obtained by fitting GreenLab outputs for the biomass of plant parts (lamina, petiole, internode, fruit), at both organ and plant level, to corresponding destructive measurements at six dates from real plants. The dynamic source–sink ratio was calculated as the ratio between biomass production and plant demand (sum of all organ sink strength) per growth cycle, both being outputs of the model.
Key Results and Conclusions
Most sink parameters were stable over multiple planting densities and seasons. From planting, source–sink ratio increased in the vegetative stage and reached a peak after fruit-set commenced, followed by a decrease of leaf appearance rate. Fruit-set probability was correlated with the source–sink ratio after the appearance of flower buds. The relationship between fruit-set probability and the most correlated source–sink ratio could be quantified by a single regression line for both experiments. The current work paves the way to predicting dynamic fruit-set using a functional structure model.
PMCID: PMC3077983  PMID: 21183453
Tomato; Solanum lycopersicum; fruit-set probability; time step; source–sink ratio; sink strength; functional–structural plant model; inverse modelling; plant plasticity
3.  Quantifying the source–sink balance and carbohydrate content in three tomato cultivars 
Supplementary lighting is frequently applied in the winter season for crop production in greenhouses. The effect of supplementary lighting on plant growth depends on the balance between assimilate production in source leaves and the overall capacity of the plants to use assimilates. This study aims at quantifying the source–sink balance and carbohydrate content of three tomato cultivars differing in fruit size, and to investigate to what extent the source/sink ratio correlates with the potential fruit size. Cultivars Komeet (large size), Capricia (medium size), and Sunstream (small size, cherry tomato) were grown from 16 August to 21 November, at similar crop management as in commercial practice. Supplementary lighting (High Pressure Sodium lamps, photosynthetic active radiation at 1 m below lamps was 162 μmol photons m-2 s-1; maximum 10 h per day depending on solar irradiance level) was applied from 19 September onward. Source strength was estimated from total plant growth rate using periodic destructive plant harvests in combination with the crop growth model TOMSIM. Sink strength was estimated from potential fruit growth rate which was determined from non-destructively measuring the fruit growth rate at non-limiting assimilate supply, growing only one fruit on each truss. Carbohydrate content in leaves and stems were periodically determined. During the early growth stage, ‘Komeet’ and ‘Capricia’ showed sink limitation and ‘Sunstream’ was close to sink limitation. During this stage reproductive organs had hardly formed or were still small and natural irradiance was high (early September) compared to winter months. Subsequently, during the fully fruiting stage all three cultivars were strongly source-limited as indicated by the low source/sink ratio (average source/sink ratio from 50 days after planting onward was 0.17, 0.22, and 0.33 for ‘Komeet,’ ‘Capricia,’ and ‘Sunstream,’ respectively). This was further confirmed by the fact that pruning half of the fruits hardly influenced net leaf photosynthesis rates. Carbohydrate content in leaves and stems increased linearly with the source/sink ratio. We conclude that during the early growth stage under high irradiance, tomato plants are sink-limited and that the level of sink limitation differs between cultivars but it is not correlated with their potential fruit size. During the fully fruiting stage tomato plants are source-limited and the extent of source limitation of a cultivar is positively correlated with its potential fruit size.
PMCID: PMC4456573  PMID: 26097485
source–sink balance; plant development stage; carbohydrate content; quantification; tomato cultivars; Solanum lycopersicum
4.  Partitioning of 13C-photosynthate from Spur Leaves during Fruit Growth of Three Japanese Pear (Pyrus pyrifolia) Cultivars Differing in Maturation Date 
Annals of Botany  2005;95(4):685-693.
• Background and Aims In fruit crops, fruit size at harvest is an important aspect of quality. With Japanese pears (Pyrus pyrifolia), later maturing cultivars usually have larger fruits than earlier maturing cultivars. It is considered that the supply of photosynthate during fruit development is a critical determinant of size. To assess the interaction of assimilate supply and early/late maturity of cultivars and its effect on final fruit size, the pattern of carbon assimilate partitioning from spur leaves (source) to fruit and other organs (sinks) during fruit growth was investigated using three genotypes differing in maturation date.
• Methods Partitioning of photosynthate from spur leaves during fruit growth was investigated by exposure of spurs to 13CO2 and measurement of the change in 13C abundance in dry matter with time. Leaf number and leaf area per spur, fresh fruit weight, cell number and cell size of the mesocarp were measured and used to model the development of the spur leaf and fruit.
• Key Results Compared with the earlier-maturing cultivars ‘Shinsui’ and ‘Kousui’, the larger-fruited, later-maturing cultivar ‘Shinsetsu’ had a greater total leaf area per spur, greater source strength (source weight × source specific activity), with more 13C assimilated per spur and allocated to fruit, smaller loss of 13C in respiration and export over the season, and longer duration of cell division and enlargement. Histology shows that cultivar differences in final fruit size were mainly attributable to the number of cells in the mesocarp.
• Conclusions Assimilate availability during the period of cell division was crucial for early fruit growth and closely correlated with final fruit size. Early fruit growth of the earlier-maturing cultivars, but not the later-maturing ones, was severely restrained by assimilate supply rather than by sink limitation.
PMCID: PMC4246860  PMID: 15655106
13C labelling; fruit growth; sink strength; spur leaves; pear; Pyrus pyrifolia Nakai
5.  Hormonal and metabolic regulation of tomato fruit sink activity and yield under salinity 
Journal of Experimental Botany  2014;65(20):6081-6095.
Cytokinins and cell wall invertase are positive players in regulating fruit sink strength, growth, and yield under salinity as components of the same signalling cascade establishing and developing sink organs.
Salinization of water and soil has a negative impact on tomato (Solanum lycopersicum L.) productivity by reducing growth of sink organs and by inducing senescence in source leaves. It has been hypothesized that yield stability implies the maintenance or increase of sink activity in the reproductive structures, thus contributing to the transport of assimilates from the source leaves through changes in sucrolytic enzymes and their regulation by phytohormones. In this study, classical and functional physiological approaches have been integrated to study the influence of metabolic and hormonal factors on tomato fruit sink activity, growth, and yield: (i) exogenous hormones were applied to plants, and (ii) transgenic plants overexpressing the cell wall invertase (cwInv) gene CIN1 in the fruits and de novo cytokinin (CK) biosynthesis gene IPT in the roots were constructed. Although salinity reduces fruit growth, sink activity, and trans-zeatin (tZ) concentrations, it increases the ethylene precursor 1-aminocyclopropane-1-carboxylic acid (ACC) during the actively growing period (25 days after anthesis). Indeed, exogenous application of the CK analogue kinetin to salinized actively growing fruits recovered sucrolytic activities (mainly cwInv and sucrose synthase), sink strength, and fruit weight, whereas the ethylene-releasing compound ethephon had a negative effect in equivalent non-stressed fruits. Fruit yield was increased by both the constitutive expression of CIN1 in the fruits (up to 4-fold) or IPT in the root (up to 30%), owing to an increase in the fruit number (lower flower abortion) and in fruit weight. This is possibly related to a recovery of sink activity in reproductive tissues due to both (i) increase in sucrolytic activities (cwInv, sucrose synthase, and vacuolar and cytoplasmic invertases) and tZ concentration, and (ii) a decrease in the ACC levels and the activity of the invertase inhibitor. This study provides new functional evidences about the role of metabolic and hormonal inter-regulation of local sink processes in controlling tomato fruit sink activity, growth, and yield under salinity.
PMCID: PMC4203140  PMID: 25170099
Cell wall invertase; cytokinins; fruit; salinity; sink activity; tomato.
6.  Dry matter partitioning models for the simulation of individual fruit growth in greenhouse cucumber canopies 
Annals of Botany  2011;108(6):1075-1084.
Background and Aims
Growth imbalances between individual fruits are common in indeterminate plants such as cucumber (Cucumis sativus). In this species, these imbalances can be related to differences in two growth characteristics, fruit growth duration until reaching a given size and fruit abortion. Both are related to distribution, and environmental factors as well as canopy architecture play a key role in their differentiation. Furthermore, events leading to a fruit reaching its harvestable size before or simultaneously with a prior fruit can be observed. Functional–structural plant models (FSPMs) allow for interactions between environmental factors, canopy architecture and physiological processes. Here, we tested hypotheses which account for these interactions by introducing dominance and abortion thresholds for the partitioning of assimilates between growing fruits.
Using the L-System formalism, an FSPM was developed which combined a model for architectural development, a biochemical model of photosynthesis and a model for assimilate partitioning, the last including a fruit growth model based on a size-related potential growth rate (RP). Starting from a distribution proportional to RP, the model was extended by including abortion and dominance. Abortion was related to source strength and dominance to sink strength. Both thresholds were varied to test their influence on fruit growth characteristics. Simulations were conducted for a dense row and a sparse isometric canopy.
Key Results
The simple partitioning models failed to simulate individual fruit growth realistically. The introduction of abortion and dominance thresholds gave the best results. Simulations of fruit growth durations and abortion rates were in line with measurements, and events in which a fruit was harvestable earlier than an older fruit were reproduced.
Dominance and abortion events need to be considered when simulating typical fruit growth traits. By integrating environmental factors, the FSPM can be a valuable tool to analyse and improve existing knowledge about the dynamics of assimilates partitioning.
PMCID: PMC3189842  PMID: 21715366
Modelling; individual fruit growth; functional–structural plant model; L-System; Cucumis sativus; cucumber; plant architecture; assimilate distribution
7.  Changes in Intake of Fruits and Vegetables and Weight Change in United States Men and Women Followed for Up to 24 Years: Analysis from Three Prospective Cohort Studies 
PLoS Medicine  2015;12(9):e1001878.
Current dietary guidelines recommend eating a variety of fruits and vegetables. However, based on nutrient composition, some particular fruits and vegetables may be more or less beneficial for maintaining or achieving a healthy weight. We hypothesized that greater consumption of fruits and vegetables with a higher fiber content or lower glycemic load would be more strongly associated with a healthy weight.
Methods and Findings
We examined the association between change in intake of specific fruits and vegetables and change in weight in three large, prospective cohorts of 133,468 United States men and women. From 1986 to 2010, these associations were examined within multiple 4-y time intervals, adjusting for simultaneous changes in other lifestyle factors, including other aspects of diet, smoking status, and physical activity. Results were combined using a random effects meta-analysis. Increased intake of fruits was inversely associated with 4-y weight change: total fruits -0.53 lb per daily serving (95% CI -0.61, -0.44), berries -1.11 lb (95% CI -1.45, -0.78), and apples/pears -1.24 lb (95% CI -1.62, -0.86). Increased intake of several vegetables was also inversely associated with weight change: total vegetables -0.25 lb per daily serving (95% CI -0.35, -0.14), tofu/soy -2.47 lb (95% CI, -3.09 to -1.85 lb) and cauliflower -1.37 lb (95% CI -2.27, -0.47). On the other hand, increased intake of starchy vegetables, including corn, peas, and potatoes, was associated with weight gain. Vegetables having both higher fiber and lower glycemic load were more strongly inversely associated with weight change compared with lower-fiber, higher-glycemic-load vegetables (p < 0.0001). Despite the measurement of key confounders in our analyses, the potential for residual confounding cannot be ruled out, and although our food frequency questionnaire specified portion size, the assessment of diet using any method will have measurement error.
Increased consumption of fruits and non-starchy vegetables is inversely associated with weight change, with important differences by type suggesting that other characteristics of these foods influence the magnitude of their association with weight change.
Using longitudinal data from health practitioners, Bertoia and colleagues explore associations between specific food choices and weight change.
Editors' Summary
Obesity—having an unhealthy amount of body fat—is increasing worldwide. In the United States, for example, more than a third of adults are obese and another third are overweight. Obesity is defined as having a body mass index (BMI; an indicator of body fat calculated by dividing a person’s weight in kilograms by their height in meters squared) of more than 30 kg/m2; overweight individuals have a BMI of 25.0–29.9 kg/m2. Compared to people with a healthy weight, overweight and obese individuals have an increased risk of developing diabetes and cardiovascular diseases (conditions that affect the heart and/or the blood vessels), and tend to die younger. People gain too much fat by consuming food and drink that contains more energy (calories) than they need for their daily activities. So, people can avoid becoming obese or reduce their BMI by eating a healthy diet that contains fewer calories and by exercising more.
Why Was This Study Done?
The 2010 Dietary Guidelines for Americans recommend that adults and children should eat a variety of fruits and vegetables to help them achieve and maintain a healthy weight. But are all fruits and vegetables equally good at controlling weight? Fruits and vegetables differ in their dietary fiber content and their glycemic load. High fiber foods increase satiety (feeling full after eating), which can reduce total energy intake. Foods with a low glycemic load produce smaller and fewer blood sugar spikes after they are consumed, which may reduce hunger later on. In this study, the researchers investigate whether consumption of fruits and vegetable with a higher fiber content or lower glycemic load is more strongly associated with a healthy weight than consumption of fruits and vegetables with a lower fiber content or higher glycemic load by analyzing data on weight and diet changes among US men and women enrolled in three large prospective cohort studies set up to examine risk factors for major chronic diseases.
What Did the Researchers Do and Find?
The researchers examined associations between changes in the intake of specific fruits and vegetables recorded in dietary questionnaires completed every 4 y and self-reported weight changes in 133,468 US men and women followed for up to 24 y. After adjusting for self-reported changes in other lifestyle factors likely to affect weight, such as smoking status and physical activity, an increased intake of fruits and of several vegetables was inversely associated with 4-y weight change. Thus, an increase in total fruit intake was associated with a change in weight over a 4-y interval of -0.53 lb (a weight loss of 0.24 kg) for each extra daily serving, and an increase in total vegetable intake was associated with a weight change of -0.25 lb (-0.11 kg) for each extra daily serving. However, increased intake of starchy vegetables such as corn, peas, and potatoes was associated with weight gain. Notably, higher-fiber, lower-glycemic load vegetables (for example, broccoli and Brussels sprouts) were more strongly inversely associated with weight change than lower-fiber, higher-glycemic load vegetables (for example, carrots and cabbage).
What Do These Findings Mean?
These findings suggest that increased consumption of fruits and non-starchy vegetables is inversely associated with weight change and that different fruits and vegetables have different effects on weight. The benefits of increased consumption were greater for fruits than for vegetables and strongest for berries, apples/pears, tofu/soy, cauliflower, and cruciferous and green leafy vegetables. Increased satiety with fewer calories could be partly responsible for the beneficial effects of increasing fruit and vegetable intake. These findings may not be generalizable—nearly all the participants were well-educated white adults. Moreover, the use of dietary questionnaires and self-reported weight measurement may have introduced measurement errors into this study and, although the researchers accounted for some key lifestyle factors that are likely to affect weight, individuals who increased their fruit and vegetable intake and lost weight may have shared other unknown characteristics that were actually responsible for their weight loss. Overall, however, these findings provide new food-specific guidance for the prevention of obesity, a primary risk factor for many life-shortening health conditions.
Additional Information
This list of resources contains links that can be accessed when viewing the PDF on a device or via the online version of the article at The World Health Organization provides information on obesity (in several languages)The Global Burden of Disease websitey provides the latest details about global obesity trends; the International Obesity Taskforce also provides information about the global obesity epidemicThe United Kingdom National Health Service Choices website provides information about obesity, cardiovascular disease, and diabetes (including some personal stories), and about healthy eatingThe American Heart Association provides information on cardiovascular disease and diabetes and on keeping healthy, including nutritional informationThe US Centers for Disease Control and Prevention has information on all aspects of overweight and is a resource provided by the US Department of Agriculture that provides individuals and healthcare professionals with user-friendly nutritional information; the 2010 Dietary Guidelines for Americans are availableMedlinePlus provides links to other sources of information on obesity, heart disease, vascular disease, and diabetes (in English and Spanish)More information about the three cohort studies that provided data for this analysis (Nurses' Health Study and Nurses' Health Study II and Health Professionals Follow-up Study) is available
PMCID: PMC4578962  PMID: 26394033
8.  Rhythms and Alternating Patterns in Plants as Emergent Properties of a Model of Interaction between Development and Functioning 
Annals of Botany  2007;101(8):1233-1242.
Background and Aims
To model plasticity of plants in their environment, a new version of the functional–structural model GREENLAB has been developed with full interactions between architecture and functioning. Emergent properties of this model were revealed by simulations, in particular the automatic generation of rhythms in plant development. Such behaviour can be observed in natural phenomena such as the appearance of fruit (cucumber or capsicum plants, for example) or branch formation in trees.
In the model, a single variable, the source–sink ratio controls different events in plant architecture. In particular, the number of fruits and branch formation are determined as increasing functions of this ratio. For some sets of well-chosen parameters of the model, the dynamical evolution of the ratio during plant growth generates rhythms.
Key Results and Conclusions
Cyclic patterns in branch formation or fruit appearance emerge without being forced by the model. The model is based on the theory of discrete dynamical systems. The mathematical formalism helps us to explain rhythm generation and to control the behaviour of the system. Rhythms can appear during both the exponential and stabilized phases of growth, but the causes are different as shown by an analytical study of the system. Simulated plant behaviours are very close to those observed on real plants. With a small number of parameters, the model gives very interesting results from a qualitative point of view. It will soon be subjected to experimental data to estimate the model parameters.
PMCID: PMC2710268  PMID: 17715304
Rhythms; plasticity; plant growth model; GREENLAB; interactions; branching system; fructification; emergent properties
9.  Tensiometer-Based Irrigation Management of Subirrigated Soilless Tomato: Effects of Substrate Matric Potential Control on Crop Performance 
Automatic irrigation scheduling based on real-time measurement of soilless substrate water status has been recognized as a promising approach for efficient greenhouse irrigation management. Identification of proper irrigation set points is crucial for optimal crop performance, both in terms of yield and quality, and optimal use of water resources. The objective of the present study was to determine the effects of irrigation management based on matric potential control on growth, plant–water relations, yield, fruit quality traits, and water-use efficiency of subirrigated (through bench system) soilless tomato. Tensiometers were used for automatic irrigation control. Two cultivars, “Kabiria” (cocktail type) and “Diana” (intermediate type), and substrate water potential set-points (−30 and −60 hPa, for “Diana,” and −30, −60, and −90 hPa for “Kabiria”), were compared. Compared with −30 hPa, water stress (corresponding to a −60 hPa irrigation set-point) reduced water consumption (14%), leaf area (18%), specific leaf area (19%), total yield (10%), and mean fruit weight (13%), irrespective of the cultivars. At −60 hPa, leaf-water status of plants, irrespective of the cultivars, showed an osmotic adjustment corresponding to a 9% average osmotic potential decrease. Total yield, mean fruit weight, plant water, and osmotic potential decreased linearly when −30, −60, and −90 hPa irrigation set-points were used in “Kabiria.” Unmarketable yield in “Diana” increased when water stress was imposed (187 vs. 349 g·plant−1, respectively, at −30 and −60 hPa), whereas the opposite effect was observed in “Kabiria,” where marketable yield loss decreased linearly [by 1.05 g·plant−1 per unit of substrate water potential (in the tested range from −30 to −90 hPa)]. In the second cluster, total soluble solids of the fruit and dry matter increased irrespective of the cultivars. In the seventh cluster, in “Diana,” only a slight increase was observed from −30 vs. −60 hPa (3.3 and 1.3%, respectively, for TSS and dry matter), whereas in “Kabiria,” the increase was more pronounced (8.7 and 12.0%, respectively, for TSS and dry matter), and further reduction in matric potential from −60 to −90 hPa confirmed the linear increase for both parameters. Both glucose and fructose concentrations increased linearly in “Kabiria” fruits on decreasing the substrate matric potential, whereas in “Diana,” there was no increase. It is feasible to act on matric potential irrigation set-points to control plant response in terms of fruit quality parameters. Precise control of substrate water status may offer the possibility to steer crop response by enhancing different crop-performance components, namely yield and fruit quality, in subirrigated tomato. Small-sized fruit varieties benefit more from controlled water stress in terms of reduced unmarketable yield loss and fruit quality improvements.
PMCID: PMC4688453  PMID: 26779189
irrigation set-point; water relations; through bench system; WUE
10.  Night temperature and source–sink effects on overall growth, cell number and cell size in bell pepper ovaries 
Annals of Botany  2012;110(5):987-994.
Background and Aims
Ovary swelling, and resultant fruit malformation, in bell pepper flowers is favoured by low night temperature or a high source–sink ratio. However, the interaction between night temperature and source–sink ratio on ovary swelling and the contribution of cell size and cell number to ovary swelling are unknown. The present research examined the interactive effects of night temperature and source–sink ratio on ovary size, cell number and cell size at anthesis in bell pepper flowers.
Bell pepper plants were grown in growth chambers at night temperatures of either 20 °C (HNT) or 12 °C (LNT). Within each temperature treatment, plants bore either 0 (non-fruiting) or two developing fruits per plant. Ovary fresh weight, cell size and cell number were measured.
Key Results
Ovary fresh weights in non-fruiting plants grown at LNT were the largest, while fresh weights were smallest in plants grown at HNT with fruits. In general, mesocarp cell size in ovaries was largest in non-fruiting plants grown at either LNT or HNT and smallest in fruiting plants at HNT. Mesocarp cell number was greater in non-fruiting plants under LNT than in the rest of the night temperature/fruiting treatments. These responses were more marked in ovaries sampled after 18 d of treatment compared with those sampled after 40 d of treatment.
Ovary fresh weight of flowers at anthesis increased 65 % in non-fruiting plants grown under LNT compared with fruiting plants grown under HNT. This increase was due primarily to increases in mesocarp cell number and size. These results indicate that the combined effects of LNT and high source–sink ratio on ovary swelling are additive. Furthermore, the combined effects of LNT and low source–sink ratio or HNT and high source–sink ratio can partially overcome the detrimental effects of LNT and high source–sink ratio.
PMCID: PMC3448427  PMID: 22933415
Capsicum annuum; fruit quality; pepper ovary swelling; source–sink effects
11.  Peach Water Relations, Gas Exchange, Growth and Shoot Mortality under Water Deficit in Semi-Arid Weather Conditions 
PLoS ONE  2015;10(4):e0120246.
In this study the sensitivity of peach tree (Prunus persica L.) to three water stress levels from mid-pit hardening until harvest was assessed. Seasonal patterns of shoot and fruit growth, gas exchange (leaf photosynthesis, stomatal conductance and transpiration) as well as carbon (C) storage/mobilization were evaluated in relation to plant water status. A simple C balance model was also developed to investigate sink-source relationship in relation to plant water status at the tree level. The C source was estimated through the leaf area dynamics and leaf photosynthesis rate along the season. The C sink was estimated for maintenance respiration and growth of shoots and fruits. Water stress significantly reduced gas exchange, and fruit, and shoot growth, but increased fruit dry matter concentration. Growth was more affected by water deficit than photosynthesis, and shoot growth was more sensitive to water deficit than fruit growth. Reduction of shoot growth was associated with a decrease of shoot elongation, emergence, and high shoot mortality. Water scarcity affected tree C assimilation due to two interacting factors: (i) reduction in leaf photosynthesis (-23% and -50% under moderate (MS) and severe (SS) water stress compared to low (LS) stress during growth season) and (ii) reduction in total leaf area (-57% and -79% under MS and SS compared to LS at harvest). Our field data analysis suggested a Ψstem threshold of -1.5 MPa below which daily net C gain became negative, i.e. C assimilation became lower than C needed for respiration and growth. Negative C balance under MS and SS associated with decline of trunk carbohydrate reserves – may have led to drought-induced vegetative mortality.
PMCID: PMC4382189  PMID: 25830350
12.  Phenology, growth and physiological adjustments of oil palm (Elaeis guineensis) to sink limitation induced by fruit pruning 
Annals of Botany  2009;104(6):1183-1194.
Background and Aims
Despite its simple architecture and small phenotypic plasticity, oil palm has complex phenology and source–sink interactions. Phytomers appear in regular succession but their development takes years, involving long lag periods between environmental influences and their effects on sinks. Plant adjustments to resulting source–sink imbalances are poorly understood. This study investigated oil palm adjustments to imbalances caused by severe fruit pruning.
An experiment with two treatments (control and complete fruit pruning) during 22 months in 2006–2008) and six replications per treatment was conducted in Indonesia. Phenology, growth of above-ground vegetative and reproductive organs, leaf morphology, inflorescence sex differentiation, dynamics of non-structural carbohydrate reserves and light-saturated net photosynthesis (Amax) were monitored.
Key Results
Artificial sink limitation by complete fruit pruning accelerated development rate, resulting in higher phytomer, leaf and inflorescence numbers. Leaf size and morphology remained unchanged. Complete fruit pruning also suppressed the abortion of male inflorescences, estimated to be triggered at about 16 months before bunch maturity. The number of female inflorescences increased after an estimated lag of 24–26 months, corresponding to time from sex differentiation to bunch maturity. The most important adjustment process was increased assimilate storage in the stem, attaining nearly 50 % of dry weight in the stem top, mainly as starch, whereas glucose, which in controls was the most abundant non-structural carbohydrate stored in oil palm, decreased.
The development rate of oil palm is in part controlled by source–sink relationships. Although increased rate of development and proportion of female inflorescences constituted observed adjustments to sink limitation, the low plasticity of plant architecture (constant leaf size, absence of branching) limited compensatory growth. Non-structural carbohydrate storage was thus the main adjustment process.
PMCID: PMC2766206  PMID: 19748908
Carbon allocation; non-structural carbohydrates; source–sink relationships; Elaeis guineensis; phenotypic plasticity; photosynthesis
13.  Competition for Assimilates and Fruit Position Affect Fruit Set in Indeterminate Greenhouse Tomato 
Annals of Botany  1995;75(1):55-65.
Localization and characterization of fruit set in winter tomato crops was investigated to determine the main internal and external controlling factors and to establish a quantitative relationship between fruit set and competition for assimilates. Individual fruit growth and development was assessed on a beef tomato cultivar during the reproductive period (first nine inflorescences). A non-destructive photograph technique was used to measure fruit growth from very early stages of their development and then calliper measurements were made on big fruits. From these measurements we determined the precise developmental stage at which fruit growth stopped. Fruit potential growth, which is defined as the growth achieved in non-limiting conditions for assimilate supply, was also assessed by this method on plants thinned to one flower per inflorescence. The latter was used to calculate the ratio between actual and potential growth, which was found to be a good index of the competition for assimilates.
Time lags of fruit set were observed mainly on distal organs. When more than three flowers were left on each inflorescence, distal organs developed at the same time as proximal organs of the following inflorescence. Consequently they were submitted to a double competition within one inflorescence and among inflorescences. It was shown that, what is commonly named ‘fruit set failure’, is not an irreversible death of the organ and that a small fruit could resume growth after a delay of several weeks as soon as the first fruits ripened and thus ceased to compete for assimilates. In that case proximal fruits resumed growth before distal ones. The delayed fruits contained only few seeds but a germination test confirmed that fertilization took place before fruit set failed.
Competition for assimilates was calculated during plant development by the ratio between actual and potential fruit growth. Potential growth of proximal fruits was strongly dependent on the position of the inflorescence on the stem, whereas potential growth of distal fruits was lower than or equal to that of proximal fruits of the same inflorescence and it was independent on the inflorescence position. We took into account both inflorescence and fruit positions to establish a quantitative relationship between fruit set of individual inflorescences and the ratio between actual and potential fruit growth.
PMCID: PMC3023665  PMID: 21247913
Tomato; Lycopersicon esculentum Mill.; fruit set; competition for assimilates; potential growth; fruit sink strength
14.  Carbon allocation during fruiting in Rubus chamaemorus 
Annals of Botany  2009;104(4):703-713.
Background and Aims
Rubus chamaemorus (cloudberry) is a herbaceous clonal peatland plant that produces an extensive underground rhizome system with distant ramets. Most of these ramets are non-floral. The main objectives of this study were to determine: (a) if plant growth was source limited in cloudberry; (b) if the non-floral ramets translocated carbon (C) to the fruit; and (c) if there was competition between fruit, leaves and rhizomes for C during fruit development.
Floral and non-floral ramet activities were monitored during the period of flower and fruit development using three approaches: gas exchange measurements, 14CO2 labelling and dry mass accumulation in the different organs. Source and sink activity were manipulated by eliminating leaves or flowers or by reducing rhizome length.
Key Results
Photosynthetic rates were lower in floral than in deflowered ramets. Autoradiographs and 14C labelling data clearly indicated that fruit is a very strong sink for the floral ramet, whereas non-floral ramets translocated C toward the rhizome but not toward floral ramets. Nevertheless, rhizomes received some C from the floral ramet throughout the fruiting period. Ramets with shorter rhizomes produced smaller leaves and smaller fruits, and defoliated ramets produced very small fruits.
Plant growth appears to be source-limited in cloudberry since a reduction in sink strength did not induce a reduction in photosynthetic activity. Non-floral ramets did not participate directly to fruit development. Developing leaves appear to compete with the developing fruit but the intensity of this competition could vary with the specific timing of the two organs. The rhizome appears to act both as a source but also potentially as a sink during fruit development. Further studies are needed to characterize better the complex role played by the rhizome in fruit C nutrition.
PMCID: PMC2729635  PMID: 19520701
Allocation pattern; 14C labelling; carbon translocation; carbon reserves; cloudberry; defoliation; fruit production; gas exchange; Rubus chamaemorus; source–sink relationship; flowering
15.  Linking ascorbic acid production in Ribes nigrum with fruit development and changes in sources and sinks 
Annals of Botany  2013;111(4):703-712.
Background and Aims
Understanding the synthesis of ascorbic acid (l-AsA) in green tissues in model species has advanced considerably; here we focus on its production and accumulation in fruit. In particular, our aim is to understand the links between organs which may be sources of l-AsA (leaves) and those which accumulate it (fruits). The work presented here tests the idea that changes in leaf and fruit number influence the accumulation of l-AsA. The aim was to understand the importance of leaf tissue in the production of l-AsA and to determine how this might provide routes for the manipulation of fruit tissue l-AsA.
The experiments used Ribes nigrum (blackcurrant), predominantly in field experiments, where the source–sink relationship was manipulated to alter potential leaf l-AsA production and fruit growth and accumulation of l-AsA. These manipulations included reductions in reproductive capacity, by raceme removal, and the availability of assimilates by leaf removal and branch phloem girdling. Natural variation in fruit growth and fruit abscission is also described as this influences subsequent experimental design and the interpretation of l-AsA data.
Key Results
Results show that fruit l-AsA concentration is conserved but total yield of l-AsA per plant is dependent on a number of innate factors many of which relate to raceme attributes. Leaf removal and phloem girdling reduced fruit weight, and a combination of both reduced fruit yields further. It appears that around 50 % of assimilates utilized for fruit growth came from apical leaves, while between 20 and 30 % came from raceme leaves, with the remainder from ‘storage’.
Despite being able to manipulate leaf area and therefore assimilate availability and stored carbohydrates, along with fruit yields, rarely were effects on fruit l-AsA concentration seen, indicating fruit l-AsA production in Ribes was not directly coupled to assimilate supply. There was no supporting evidence that l-AsA production occurred predominantly in green leaf tissue followed by its transfer to developing fruits. It is concluded that l-AsA production occurs predominantly in the fruit of Ribes nigrum.
PMCID: PMC3605959  PMID: 23419248
l-Ascorbic acid; blackcurrant; fruit; Ribes nigrum; source sinks; vitamin C
16.  Pepper Weevil (Coleoptera: Curculionidae) Preferences for Specific Pepper Cultivars, Plant Parts, Fruit Colors, Fruit Sizes, and Timing 
Insects  2016;7(1):9.
Peppers (Capsicum spp.) are an important crop in the USA, with about 32,000 ha cultivated in 2007, which resulted in $588 million in farm revenue. The pepper weevil, Anthonomus eugenii Cano (Coleoptera: Curculionidae), is the most troublesome insect pest of peppers in the southern United States. It is therefore urgent to find different vulnerabilities of pepper cultivars, fruit and plants parts, fruit colors and sizes, and timing to infestation by A. eugenii. Also relevant is testing whether fruit length and infestation state affect fruit numbers, weights, and proportions of fruit that are infested. Counts of A. eugenii adults and marks from oviposition and feeding suggested that C. chinense Jacquin “Habanero” was least susceptible, and C. annuum L. cultivars “SY” and “SR” were most susceptible. Comparison of plant parts and fruit sizes revealed that A. eugenii preferred the peduncle, calyx, and top of pepper fruits over the middle, bottom, leaves, or remainder of flowers. Anthonomus eugenii does not discriminate between green or yellow fruit color nor vary diurnally in numbers. Based on adult counts, medium to extra-large fruits (≥1.5 cm long) attracted more weevils than small fruits (<1.5 cm). However based on proportions of fruit numbers or fruit weights that were infested, there were no differences between large and small fruits. Choice of pepper cultivar can thus be an important part of an IPM cultural control program designed to combat A. eugenii by reduced susceptibility or by synchronous fruit drop of infested fruits. Our results are potentially helpful in developing scouting programs including paying particular attention to the preferred locations of adults and their sites of feeding and oviposition on the fruit. The results also suggested the potential value of spraying when the fruits are still immature to prevent and control infestation.
PMCID: PMC4808789  PMID: 26959066
Anthonomus eugenii; cultural control; scouting
17.  Regulation Effects of Water and Nitrogen on the Source-Sink Relationship in Potato during the Tuber Bulking Stage 
PLoS ONE  2016;11(1):e0146877.
The source-sink relationship determines crop yield, and it is largely regulated by water and nutrients in agricultural production. This has been widely investigated in cereals, but fewer studies have been conducted in root and tuber crops such as potato (Solanum tuberosum L.). The objective of this study was to investigate the source-sink relationship in potato and the regulation of water and nitrogen on the source-sink relationship during the tuber bulking stage. A pot experiment using virus-free plantlets of the Atlantic potato cultivar was conducted, using three water levels (50%, 70% and 90% of field capacity) and three nitrogen levels (0, 0.2, 0.4 g N∙kg−1 soil). The results showed that, under all water and nitrogen levels, plant source capacity were small at the end of the experiment, since photosynthetic activity in leaves were low and non-structural reserves in underground stems were completely remobilized. While at this time, there were very big differences in maximum and minimum tuber number and tuber weight, indicating that the sink tuber still had a large potential capacity to take in assimilates. These results suggest that the source-supplied assimilates were not sufficient enough to meet the demands of sink growth. Thus, we concluded that, unlike cereals, potato yield is more likely to be source-limited than sink-limited during the tuber bulking stage. Water and nitrogen are two key factors in potato production management. Our results showed that water level, nitrogen level and the interaction between water and nitrogen influence potato yield mainly through affecting source capacity via the net photosynthetic rate, total leaf area and leaf life span. Well-watered, sufficient nitrogen and well-watered combined with sufficient nitrogen increased yield mainly by enhancing the source capacity. Therefore, this suggests that increasing source capacity is more crucial to improve potato yield.
PMCID: PMC4709089  PMID: 26752657
18.  Metabolomic profiling in tomato reveals diel compositional changes in fruit affected by source–sink relationships 
Journal of Experimental Botany  2015;66(11):3391-3404.
Using metabolomics in tomato, we confirmed the existence of diel patterns in leaf composition and showed lower but significant diel changes in expanding fruit depending on the potential carbon supply.
A detailed study of the diurnal compositional changes was performed in tomato (Solanum lycopersicum cv. Moneymaker) leaves and fruits. Plants were cultivated in a commercial greenhouse under two growth conditions: control and shaded. Expanding fruits and the closest mature leaves were harvested during two different day/night cycles (cloudy or sunny day). High-throughput robotized biochemical phenotyping of major compounds, as well as proton nuclear magnetic resonance and mass spectrometry metabolomic profiling, were used to measure the contents of about 70 metabolites in the leaves and 60 metabolites in the fruits, in parallel with ecophysiological measurements. Metabolite data were processed using multivariate, univariate, or clustering analyses and correlation networks. The shaded carbon-limited plants adjusted their leaf area, decreased their sink carbon demand and showed subtle compositional modifications. For source leaves, several metabolites varied along a diel cycle, including those directly linked to photosynthesis and photorespiration. These metabolites peaked at midday in both conditions and diel cycles as expected. However, transitory carbon storage was limited in tomato leaves. In fruits, fewer metabolites showed diel fluctuations, which were also of lower amplitude. Several organic acids were among the fluctuating metabolites. Diel patterns observed in leaves and especially in fruits differed between the cloudy and sunny days, and between the two conditions. Relationships between compositional changes in leaves and fruits are in agreement with the fact that several metabolic processes of the fruit appeared linked to its momentary supply of sucrose.
PMCID: PMC4449552  PMID: 25873655
Diurnal changes; fruit metabolism; 1H-NMR; MS; metabolomics; Solanum lycopersicum.
19.  Trade-Offs of Flowering and Maturity Synchronisation for Pineapple Quality 
PLoS ONE  2015;10(11):e0143290.
In the pineapple sector of Benin, poor fruit quality prevents pineapple producers to enter the European market. We investigated effects of common cultural practices, flowering and maturity synchronisation, (1) to quantify the trade-offs of flowering and maturity synchronisation for pineapple quality and the proportion of fruits exportable to European markets, and (2) to determine the effect of harvesting practice on quality attributes. Four on-farm experiments were conducted during three years using cultivars Sugarloaf and Smooth Cayenne. A split-split plot design was used in each experiment, with flowering induction practice as main factor (artificial or natural flowering induction), maturity induction practice as split factor (artificial or natural maturity induction) and harvesting practice as the split-split factor (farmers’ harvest practice or individual fruit harvesting at optimum maturity). Artificial flowering induction gave fruits with lower infructescence weight, higher ratio crown: infructescence length, and a lower proportion of fruits exportable to European markets than natural flowering induction. The costs of the improvements by natural flowering induction were huge: the longer durations from planting to flowering induction and harvesting, the higher number of harvestings of the fruits increasing the labour cost and the lower proportion of plants producing fruits compared with crops from artificially flowering-induced plants. Artificial maturity induction decreased the total soluble solids concentration in the fruits compared with natural maturity induction thus decreasing the proportion of fruits exportable to European markets, at a benefit of only a slightly shorter time from flowering induction to harvesting. Harvesting individual fruits at optimum maturity gave fruits with higher total soluble solids in naturally maturity induced fruits compared with the farmers’ harvest practice. Given the huge costs of natural flowering induction, options to use artificial flowering induction effectively for obtaining high fruit quality are discussed.
PMCID: PMC4657890  PMID: 26599545
20.  Parameter Optimization and Field Validation of the Functional–Structural Model GREENLAB for Maize at Different Population Densities 
Annals of Botany  2007;101(8):1185-1194.
Background and Aims
Plant population density (PPD) influences plant growth greatly. Functional–structural plant models such as GREENLAB can be used to simulate plant development and growth and PPD effects on plant functioning and architectural behaviour can be investigated. This study aims to evaluate the ability of GREENLAB to predict maize growth and development at different PPDs.
Two field experiments were conducted on irrigated fields in the North China Plain with a block design of four replications. Each experiment included three PPDs: 2·8, 5·6 and 11·1 plants m−2. Detailed observations were made on the dimensions and fresh biomass of above-ground plant organs for each phytomer throughout the seasons. Growth stage-specific target files (a description of plant organ weight and dimension according to plant topological structure) were established from the measured data required for GREENLAB parameterization. Parameter optimization was conducted using a generalized least square method for the entire growth cycles for all PPDs and years. Data from in situ plant digitization were used to establish geometrical symbol files for organs that were then applied to translate model output directly into 3-D representation for each time step of the model execution.
Key Results
The analysis indicated that the parameter values of organ sink variation function, and the values of most of the relative sink strength parameters varied little among years and PPDs, but the biomass production parameter, computed plant projection surface and internode relative sink strength varied with PPD. Simulations of maize plant growth based on the fitted parameters were reasonably good as indicated by the linearity and slopes similar to unity for the comparison of simulated and observed values. Based on the parameter values fitted from different PPDs, shoot (including vegetative and reproductive parts of the plant) and cob fresh biomass for other PPDs were simulated. Three-dimensional representation of individual plant and plant stand from the model output with two contrasting PPDs were presented with which the PPD effect on plant growth can be easily recognized.
This study showed that GREENLAB model has the ability to capture plant plasticity induced by PPD. The relatively stable parameter values strengthened the hypothesis that one set of equations can govern dynamic organ growth. With further validation, this model can be used for agronomic applications such as yield optimization.
PMCID: PMC2710275  PMID: 17921525
Functional–structural plant model; GREENLAB; plant architecture; source–sink relationship; plant population density; maize (Zea mays); model parameterization
21.  Olive Fertility as Affected by Cross-Pollination and Boron 
The Scientific World Journal  2012;2012:375631.
Self-compatibility of local olive (Olea europaea L.) accessions and of the cultivars “Frantoio” and “Leccino” was investigated in Garda Lake area, northern Italy. Intercompatibility was determined for “Casaliva,” “Frantoio,” and “Leccino,” as well as the effects of foliar Boron applications (0, 262, 525, or 1050 mg·L−1) applied about one week before anthesis on fruit set, shotberry set, and on in vitro pollen germination. Following self-pollination, fruit set was significantly lower and the occurrence of shot berries significantly higher than those obtained by open pollination. No significant effect of controlled cross-pollination over self-pollination on fruit set and shotberry set was detectable. B treatments increased significantly fruit set in “Frantoio” and “Casaliva” but not in “Leccino.” B sprays had no effect on shotberry set, suggesting that these parthenocarpic fruits did not strongly compete for resources allocation and did not take advantage of increased B tissue levels. Foliar B application enhanced in vitro pollen germination, and the optimal level was higher for pollen germination than for fruit set. Our results highlight the importance of olive cross pollination for obtaining satisfactory fruit set and the beneficial effect of B treatments immediately prior to anthesis, possibly by affecting positively the fertilisation process and subsequent plant source-sink relations linked to fruitlet retention.
PMCID: PMC3417202  PMID: 22919310
22.  Heterogeneity in pineapple fruit quality results from plant heterogeneity at flower induction 
Heterogeneity in fruit quality constitutes a major constraint in agri-food chains. In this paper the sources of the heterogeneity in pineapple in the field were studied in four experiments in commercial pineapple fields. The aims were to determine (a) whether differences in pineapple fruit quality among individual fruits are associated with differences in vigor of the individual plants within the crop at the time of artificial flower induction; and (b) whether the side shoots produced by the plant during the generative phase account for the fruit quality heterogeneity. Two pineapple cultivars were considered: cv. Sugarloaf and cv. Smooth Cayenne. Plant vigor at the time of artificial flower induction was measured by three variates: the number of functional leaves, the D-leaf length and their cross product. Fruit quality attributes measured at harvest time included external attributes (weight and height of fruit, infructescence and crown) and internal quality attributes [total soluble solids (TSS), pH, translucent flesh]. Results showed that the heterogeneity in fruit weight was a consequence of the heterogeneity in vigor of the plants at the moment of flower induction; that effect was mainly on the infructescence weight and less or not on the crown weight. The associations between plant vigor variates at flower induction and the internal quality attributes of the fruit were poor and/or not consistent across experiments. The weight of the slips (side shoots) explained part of the heterogeneity in fruit weight, infructescence weight and fruit height in cv. Sugarloaf. Possibilities for reducing the variation in fruit quality by precise cultural practices are discussed.
PMCID: PMC4260489  PMID: 25538714
Ananas comosus; D-leaf; fruit size; variation; variation in quality; variation within crop; vigor
23.  Use of Fertigation and Municipal Solid Waste Compost for Greenhouse Pepper Cultivation 
The Scientific World Journal  2012;2012:973193.
Municipal solid waste compost (MSWC) and/or fertigation used in greenhouse pepper (Capsicum annuum L.) cultivation with five different substrates with soil (S) and/or MSWC mixtures (0–5–10–20–40%) used with or without fertigation. Plants growth increased in 10–20% MSWC and fertigation enhanced mainly the plant height. Fruit number increased in S : MSWC 80 : 20 without fertilizer. Plant biomass increased as MSWC content increased. There were no differences regarding leaf fluoresces and plant yield. The addition of MSWC increased nutritive value (N, K, P, organic matter) of the substrate resulting in increased EC. Fruit fresh weight decreased (up to 31%) as plants grown in higher MSWC content. Fruit size fluctuated when different MSWC content used into the soil and the effects were mainly in fruit diameter rather than in fruit length. Interestingly, the scale of marketable fruits reduced as MSWC content increased into the substrate but addition of fertilizer reversed this trend and maintained the fruit marketability. MSWC affected quality parameters and reduced fruit acidity, total phenols but increased fruit lightness. No differences observed in fruit dry matter content, fruit firmness, green colour, total soluble sugars and EC of peppers and bacteria (total coliform and E. coli) units. Low content of MSWC improved plant growth and maintained fruit fresh weight for greenhouse pepper without affecting plant yield, while fertigation acted beneficially.
PMCID: PMC3356757  PMID: 22645489
24.  In-silico analysis of water and carbon relations under stress conditions. A multi-scale perspective centered on fruit 
Fruit development, from its early stages, is the result of a complex network of interacting processes, on different scales. These include cell division, cell expansion but also nutrient transport from the plant, and exchanges with the environment. In the presence of nutrient limitation, in particular, the plant reacts as a whole, by modifying its architecture, metabolism, and reproductive strategy, determining the resources available for fruit development, which in turn affects the overall source-sink balance of the system. Here, we present an integrated model of tomato that explicitly accounts for early developmental changes (from cell division to harvest), and use it to investigate the impact of water deficit and carbon limitation on nutrient fluxes and fruit growth, in both dry and fresh mass. Variability in fruit response is analyzed on two different scales: among trusses at plant level, and within cell populations at fruit level. Results show that the effect of stress on individual cells strongly depends on their age, size, and uptake capabilities, and that the timing of stress application, together with the fruit position on the plant, is crucial in determining the final phenotypic outcome. Water deficit and carbon depletion impacted either source size, source activity, or sink strength with contrasted effects on fruit growth. An important prediction of the model is the major role of symplasmic transport of carbon in the early stage of fruit development, as a catalyst for cell and fruit growth.
PMCID: PMC3856696  PMID: 24367372
model; development; cell division; cell expansion; symplasm; plant architecture; tomato; stress
25.  The Derivation of Sink Functions of Wheat Organs using the GREENLAB Model 
Annals of Botany  2007;101(8):1099-1108.
Background and Aims
In traditional crop growth models assimilate production and partitioning are described with empirical equations. In the GREENLAB functional–structural model, however, allocation of carbon to different kinds of organs depends on the number and relative sink strengths of growing organs present in the crop architecture. The aim of this study is to generate sink functions of wheat (Triticum aestivum) organs by calibrating the GREENLAB model using a dedicated data set, consisting of time series on the mass of individual organs (the ‘target data’).
An experiment was conducted on spring wheat (Triticum aestivum, ‘Minaret’), in a growth chamber from, 2004 to, 2005. Four harvests were made of six plants each to determine the size and mass of individual organs, including the root system, leaf blades, sheaths, internodes and ears of the main stem and different tillers. Leaf status (appearance, expansion, maturity and death) of these 24 plants was recorded. With the structures and mass of organs of four individual sample plants, the GREENLAB model was calibrated using a non-linear least-square-root fitting method, the aim of which was to minimize the difference in mass of the organs between measured data and model output, and to provide the parameter values of the model (the sink strengths of organs of each type, age and tiller order, and two empirical parameters linked to biomass production).
Key Results and Conclusions
The masses of all measured organs from one plant from each harvest were fitted simultaneously. With estimated parameters for sink and source functions, the model predicted the mass and size of individual organs at each position of the wheat structure in a mechanistic way. In addition, there was close agreement between experimentally observed and simulated values of leaf area index.
PMCID: PMC2710274  PMID: 18045794
Wheat; Triticum aestivum ‘Minaret’; tiller; GREENLAB; organ mass; functional–structural model; model calibration; multi-fitting; source–sink

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