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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.  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
4.  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.
5.  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
6.  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
7.  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
8.  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
9.  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
10.  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
11.  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
12.  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
13.  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
14.  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
15.  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
16.  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
17.  Time‐course of Tomato Whole‐plant Respiration and Fruit and Stem Growth During Prolonged Darkness in Relation to Carbohydrate Reserves 
Annals of Botany  2003;91(4):429-438.
To evaluate the relevance of a simple carbon balance model (Seginer et al., 1994, Scientia Horticulturae 60: 55–80) in source‐limiting conditions, the dynamics of growth, respiration and carbohydrate reserves of tomato plants were observed in prolonged darkness. Four days prior to the experiments, plants were exposed to high or low light levels and CO2 concentrations. The concentration of carbohydrates in vegetative organs was 30–50 % lower in plants that were exposed to low carbon assimilation conditions compared with those exposed to high carbon assimilation conditions. During prolonged darkness, plants with low carbohydrate reserves exhibited a lower whole‐plant respiration rate, which decreased rapidly to almost zero after 24 h, and carbohydrate pools were almost exhausted in leaves, roots and flowers. In plants with high carbohydrate reserves, the whole‐plant respiration rate was maintained for a longer period and carbohydrates remained available for at least 48 h in leaves and flowers. In contrast, fruits maintained fairly stable and identical concentrations of carbohydrates and the reduction in their rate of expansion was moderate irrespective of the pre‐treatment carbon assimilation conditions. The time‐course of asparagine and glutamine concentrations showed the occurrence of carbon stress in leaves and flowers. Estimation of source and sink activities indicated that even after low carbon assimilation, vegetative organs contained enough carbohydrates to support fruit growth provided their own growth stopped. The time of exhaustion of these carbohydrates corresponded grossly to the maintenance stage simulated by the model proposed by Seginer et al. (1994), thus validating the use of such a model for optimizing plant growth.
PMCID: PMC4241059  PMID: 12588722
Tomato; Lycopersicon esculentum Mill.; prolonged darkness; respiration; carbohydrate pools; fruit growth; carbon stress; source–sink balance
18.  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
19.  Critical review: vegetables and fruit in the prevention of chronic diseases 
European Journal of Nutrition  2012;51(6):637-663.
Vegetables and fruit provide a significant part of human nutrition, as they are important sources of nutrients, dietary fibre, and phytochemicals. However, it is uncertain whether the risk of certain chronic diseases can be reduced by increased consumption of vegetables or fruit by the general public, and what strength of evidence has to be allocated to such an association.
Therefore, a comprehensive analysis of the studies available in the literature and the respective study results has been performed and evaluated regarding obesity, type 2 diabetes mellitus, hypertension, coronary heart disease (CHD), stroke, cancer, chronic inflammatory bowel disease (IBD), rheumatoid arthritis (RA), chronic obstructive pulmonary disease (COPD), asthma, osteoporosis, eye diseases, and dementia. For judgement, the strength of evidence for a risk association, the level of evidence, and the number of studies were considered, the quality of the studies and their estimated relevance based on study design and size.
For hypertension, CHD, and stroke, there is convincing evidence that increasing the consumption of vegetables and fruit reduces the risk of disease. There is probable evidence that the risk of cancer in general is inversely associated with the consumption of vegetables and fruit. In addition, there is possible evidence that an increased consumption of vegetables and fruit may prevent body weight gain. As overweight is the most important risk factor for type 2 diabetes mellitus, an increased consumption of vegetables and fruit therefore might indirectly reduces the incidence of type 2 diabetes mellitus. Independent of overweight, there is probable evidence that there is no influence of increased consumption on the risk of type 2 diabetes mellitus. There is possible evidence that increasing the consumption of vegetables and fruit lowers the risk of certain eye diseases, dementia and the risk of osteoporosis. Likewise, current data on asthma, COPD, and RA indicate that an increase in vegetable and fruit consumption may contribute to the prevention of these diseases. For IBD, glaucoma, and diabetic retinopathy, there was insufficient evidence regarding an association with the consumption of vegetables and fruit.
This critical review on the associations between the intake of vegetables and fruit and the risk of several chronic diseases shows that a high daily intake of these foods promotes health. Therefore, from a scientific point of view, national campaigns to increase vegetable and fruit consumption are justified. The promotion of vegetable and fruit consumption by nutrition and health policies is a preferable strategy to decrease the burden of several chronic diseases in Western societies.
PMCID: PMC3419346  PMID: 22684631
Vegetables; Fruit; Prevention; Chronic diseases; Epidemiology
20.  Effect of heat treatment around the fruit set region on growth and yield of watermelon [Citrullus lanatus (Thunb.) Matsum. and Nakai] 
The present experiment was aimed to study the effect of imposing modulated temperature treatments 14 °C and 18 °C, around the fruiting region of watermelon plants, and to estimate the economic feasibility of the temperature treatments based on energy consumption for heating. Watermelon cultivar ‘Sambok-gul’ was selected and sown on perlite beds in a plastic house under controlled conditions at Watermelon Farm, Jeongeup-Jeonbuk, longitude 35° 31′ 47.51N, 126° 48′48.84E, altitude 37 m during the early spring season (2010–2011). The findings revealed that the temperature treatment at 18 °C caused significant increase in weight (2.0 kg plant−1), fruit weight (8.3 kg plant−1), soluble solid content (11.5 %), and fruit set rate (96.5 %) at harvest stage. Higher contents of Ca2+ and Mg2+ ions were observed in the 1st upper leaf of the fruit set node (79.3 mg L−1) and the 1st lower leaf of the fruit set node (12.0 mg L−1), respectively at 14 °C. The power consumption and extra costs of the temperature treatment 18 °C were suggested as affordable and in range of a farmer’s budget (41.14 USD/22 days). Hence, it was concluded that modulating temperature treatments could be utilized successfully to optimize the temperature range for enhancing the fruit yield and quality in the winter watermelon crops.
PMCID: PMC3781286  PMID: 24431519
Temperature; Electricity consumption; Yield; Soluble solid content; Ion content
21.  Genetic mechanisms underlying yield potential in the rice high-yielding cultivar Takanari, based on reciprocal chromosome segment substitution lines 
BMC Plant Biology  2014;14(1):295.
Increasing rice yield potential is a major objective in rice breeding programs, given the need for meeting the demands of population growth, especially in Asia. Genetic analysis using genomic information and high-yielding cultivars can facilitate understanding of the genetic mechanisms underlying rice yield potential. Chromosome segment substitution lines (CSSLs) are a powerful tool for the detection and precise mapping of quantitative trait loci (QTLs) that have both large and small effects. In addition, reciprocal CSSLs developed in both parental cultivar backgrounds may be appropriate for evaluating gene activity, as a single factor or in epistatic interactions.
We developed reciprocal CSSLs derived from a cross between Takanari (one of the most productive indica cultivars) and a leading japonica cultivar, Koshihikari; both the cultivars were developed in Japan. Forty-one CSSLs covered most of the Takanari genome in the Koshihikari background and 39 CSSLs covered the Koshihikari genome in the Takanari background. Using the reciprocal CSSLs, we conducted yield trials under canopy conditions in paddy fields. While no CSSLs significantly exceeded the recurrent parent cultivar in yield, genetic analysis detected 48 and 47 QTLs for yield and its components in the Koshihikari and Takanari backgrounds, respectively. A number of QTLs showed a trade-off, in which the allele with increased sink-size traits (spikelet number per panicle or per square meter) was associated with decreased ripening percentage or 1000-grain weight. These results indicate that increased sink size is not sufficient to increase rice yield in both backgrounds. In addition, most QTLs were detected in either one of the two genetic backgrounds, suggesting that these loci may be under epistatic control with other gene(s).
We demonstrated that the reciprocal CSSLs are a useful tool for understanding the genetic mechanisms underlying yield potential in the high-yielding rice cultivar Takanari. Our results suggest that sink-size QTLs in combination with QTLs for source strength or translocation capacity, as well as careful attention to epistatic interactions, are necessary for increasing rice yield. Thus, our findings provide a foundation for developing rice cultivars with higher yield potential in future breeding programs.
Electronic supplementary material
The online version of this article (doi:10.1186/s12870-014-0295-2) contains supplementary material, which is available to authorized users.
PMCID: PMC4243286  PMID: 25404368
Chromosome segment substitution lines (CSSLs); Quantitative trait locus (QTL); Rice; Yield potential
22.  Tomato fruits: a good target for iodine biofortification 
Iodine is a trace element that is fundamental for human health: its deficiency affects about two billion people worldwide. Fruits and vegetables are usually poor sources of iodine; however, plants can accumulate iodine if it is either present or exogenously administered to the soil. The biofortification of crops with iodine has therefore been proposed as a strategy for improving human nutrition. A greenhouse pot experiment was carried out to evaluate the possibility of biofortifying tomato fruits with iodine. Increasing concentrations of iodine supplied as KI or KIO3 were administered to plants as root treatments and the iodine accumulation in fruits was measured. The influences of the soil organic matter content or the nitrate level in the nutritive solution were analyzed. Finally, yield and qualitative properties of the biofortified tomatoes were considered, as well as the possible influence of fruit storage and processing on the iodine content. Results showed that the use of both the iodized salts induced a significant increase in the fruit’s iodine content in doses that did not affect plant growth and development. The final levels ranged from a few mg up to 10 mg iodine kg - 1 fruit fresh weight and are more than adequate for a biofortification program, since 150 μg iodine per day is the recommended dietary allowance for adults. In general, the iodine treatments scarcely affected fruit appearance and quality, even with the highest concentrations applied. In contrast, the use of KI in plants fertilized with low doses of nitrate induced moderate phytotoxicity symptoms. Organic matter-rich soils improved the plant’s health and production, with only mild reductions in iodine stored in the fruits. Finally, a short period of storage at room temperature or a 30-min boiling treatment did not reduce the iodine content in the fruits, if the peel was maintained. All these results suggest that tomato is a particularly suitable crop for iodine biofortification programs.
PMCID: PMC3694224  PMID: 23818889
biofortification; iodine; iodine deficiency; potassium iodate; potassium iodide; Solanum lycopersicum L.; tomato
23.  A Thirty-Year Survey Reveals That Ecosystem Function of Fungi Predicts Phenology of Mushroom Fruiting 
PLoS ONE  2012;7(11):e49777.
Mushroom fruiting, the reproduction of fungi, has broad implications for forest health, terrestrial biomass turnover, and global carbon cycle. However, little is known about the difference in phenology and environmental drivers of mushroom fruiting between functional guilds, e.g., ectomycorrhizal (ECM) mutualists and saprotrophs (SAP). There is a remarkable difference between ECM and SAP fungi in their available carbon sources and lifecycles, and thus these fungal groups are likely to differ in fruiting phenology. We analyzed intra- and inter-annual phenological patterns of mushroom fruiting throughout the year using a long-term census dataset of mushroom-forming fungi in a Japanese oak forest in which a total of 11,923 mushroom counts (668 species) were recorded during monthly intervals from 1982 to 2011. ECM fungi showed a unimodal seasonal fruiting peak from mid-summer to early autumn; litter-decomposing fungi showed moderate fruiting peaks from early summer or early autumn, and the phenology of wood-decomposing fungi varied considerably among the genera. Each functional group was controlled by a different set of external factors; temperature and rainfall increased ECM fungal fruiting, but key factors substantially differed among the genera of litter- and wood-decomposing fungi in taxon-specific ways. Our results suggest that fungal fruiting phenology may be affected by the seasonality of carbohydrate availability. The highly scheduled reproduction of ECM fungi may reflect temperature-dependent increases and drought-induced decreases of photosynthetic activity in host plants rather than improved growth conditions for fungi during the summer. We argue that the way a fungus obtains carbohydrates may explain a substantial fraction of the fruiting phenology, which may make a differential contribution to the community structure of fungus-associated organisms and terrestrial biomass turnover based on fungal functional groups.
PMCID: PMC3507881  PMID: 23209598
24.  Uptake and Distribution of Cd in Sweet Maize Grown on Contaminated Soils: A Field-Scale Study 
Maize is an economic crop that is also a candidate for use in phytoremediation in low-to-moderately Cd-contaminated soils, because the plant can accumulate high concentration of Cd in parts that are nonedible to humans while accumulating only a low concentration of Cd in the fruit. Maize cultivars CT38 and HZ were planted in field soils contaminated with Cd and nitrilotriacetic acid (NTA) was used to enhance the phytoextractive effect of the maize. Different organs of the plant were analyzed to identify the Cd sinks in the maize. A distinction was made between leaf sheath tissue and leaf lamina tissue. Cd concentrations decreased in the tissues in the following order: sheath > root > lamina > stem > fruit. The addition of NTA increased the amount of Cd absorbed but left the relative distribution of the metal among the plant organs essentially unchanged. The Cd in the fruit of maize was below the Chinese government's permitted concentration in coarse cereals. Therefore, this study shows that it is possible to conduct maize phytoremediation of Cd-contaminated soil while, at the same time, harvesting a crop, for subsequent consumption.
PMCID: PMC3856120  PMID: 24348276
25.  A functional–structural kiwifruit vine model integrating architecture, carbon dynamics and effects of the environment 
Annals of Botany  2010;107(5):747-764.
Background and Aims
Functional–structural modelling can be used to increase our understanding of how different aspects of plant structure and function interact, identify knowledge gaps and guide priorities for future experimentation. By integrating existing knowledge of the different aspects of the kiwifruit (Actinidia deliciosa) vine's architecture and physiology, our aim is to develop conceptual and mathematical hypotheses on several of the vine's features: (a) plasticity of the vine's architecture; (b) effects of organ position within the canopy on its size; (c) effects of environment and horticultural management on shoot growth, light distribution and organ size; and (d) role of carbon reserves in early shoot growth.
Using the L-system modelling platform, a functional–structural plant model of a kiwifruit vine was created that integrates architectural development, mechanistic modelling of carbon transport and allocation, and environmental and management effects on vine and fruit growth. The branching pattern was captured at the individual shoot level by modelling axillary shoot development using a discrete-time Markov chain. An existing carbon transport resistance model was extended to account for several source/sink components of individual plant elements. A quasi-Monte Carlo path-tracing algorithm was used to estimate the absorbed irradiance of each leaf.
Key Results
Several simulations were performed to illustrate the model's potential to reproduce the major features of the vine's behaviour. The model simulated vine growth responses that were qualitatively similar to those observed in experiments, including the plastic response of shoot growth to local carbon supply, the branching patterns of two Actinidia species, the effect of carbon limitation and topological distance on fruit size and the complex behaviour of sink competition for carbon.
The model is able to reproduce differences in vine and fruit growth arising from various experimental treatments. This implies it will be a valuable tool for refining our understanding of kiwifruit growth and for identifying strategies to improve production.
PMCID: PMC3077975  PMID: 20855486
Actinidia deliciosa; kiwifruit; L-systems; plant architecture; carbon allocation; functional–structural plant model

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