Effect of temperature on biomass allocation in tomato (Lycopersicon esculentum)

Temperature may influence dry matter partitioning between fruits and vegetative plant parts either directly or indirectly through its influence on development, flower and/or fruit abortion. The objective of the present work was to investigate whether there is any direct effect of temperature on dry matter partitioning between fruits and vegetative plant parts in tomato. A greenhouse experiment was conducted, with alternating 3-week periods of high (23°C) and low (18°C) temperature setpoint. Dry matter partitioning during these 3-week periods was determined from destructive plant harvests at two levels of fruit pruning (3 and 7 fruits per truss). Indirect temperature effects on dry matter partitioning were excluded by fruit pruning.
On average, the fraction of dry matter distributed to the fruits during a 12-week period, starting with the flowering of the fifth truss (28 days after planting), was 0.53 (3 fruits per truss) and 0.70 (7 fruits per truss). These ratios were also calculated for every 3-week period separately and did not depend on the average temperature (18–24°C) during that period.
It is concluded that dry matter distribution in tomato is not significantly affected by temperature directly, which means that the temperature effect (18–24°C) on the generative sink strength is not much different from the temperature effect on the vegetative sink strength.     

Developmental characterization of the fasciated locus and mapping of Arabidopsis candidate genes involved in the control of floral meristem size and carpel number in tomato.

Mutation at the fasciated locus was a key step in the production of extreme fruit size during tomato domestication. To shed light on the nature of these changes, near-isogenic lines were used for a comparative developmental study of fasciated and wild-type tomato plants. The fasciated gene directly affects floral meristem size and is expressed before the earliest stages of flower organogenesis. As a result, mature fruit of fasciated mutants have more carpels (locules) and greater fruit diameter and mass. The discovery that fasciated affects floral meristem size led to a search for candidate genes from Arabidopsis known to be involved in floral meristem development. Putative homologs were identified in a large tomato EST database, verified through phylogenetic analyses, and mapped in tomato; none mapped to the fasciated locus; however, putative homologs of WUS and WIG mapped to the locule number locus on chromosome 2, the second major transition to large tomato fruit, with WUS showing the highest association. In other cases, minor QTLs for floral organ number (lcn2.2) and (stn11.2) co-localized with a CLV1 paralog and with the syntenic region containing the CLV3 gene in Arabidopsis, respectively.     

The effect of assimilate supply on fruit growth and hormone levels in tomato plants

Rates of dry matter accumulation and contents of starch, sugars and abscisic acid (ABA) of tomato fruits differed significantly during development at three positions (proximal, middle and distal) on a truss. Proximal fruits, which accumulated dry matter most rapidly during early development, generally had least ABA (per g DW).Partial defoliation reduced carbon accumulation by all fruits but increased ABA, especially in distal fruit, and indoleacetic acid (IAA), particularly in proximal fruits. The ABA content of leaves in partially defoliated plants was similar to that of leaves on non-defoliated plants.Removal of distal fruits on a truss enhanced carbon movement to the remaining proximal fruits and also increased their ABA content early in development but did not affect their IAA content. On the other hand, when proximal fruit were removed there was no large or lasting increased accumulation of carbon by the remaining distal fruits and they contained less ABA and IAA than fruits on plants without fruit thinning. Leaf carbon and ABA levels showed no marked trend in response to fruit thinning.The amount of carbon in the stems was increased by fruit thinning but decreased by partial defoliation.The possible roles of ABA and IAA in regulating fruit growth are discussed.Part of this work has been presented to a Symposium on Phloem loading and related processes at Bad Grund/Oberhar, W. Germany, July 1979.     

Aucsia gene silencing causes parthenocarpic fruit development in tomato

In angiosperms, auxin phytohormones play a crucial regulatory role in fruit initiation. The expression of auxin biosynthesis genes in ovules and placenta results in uncoupling of tomato (Solanum lycopersicum) fruit development from fertilization with production of parthenocarpic fruits. We have identified two newly described genes, named Aucsia genes, which are differentially expressed in auxin-synthesis (DefH9-iaaM) parthenocarpic tomato flower buds. The two tomato Aucsia genes encode 53-amino-acid-long peptides. We show, by RNA interference-mediated gene suppression, that Aucsia genes are involved in both reproductive and vegetative plant development. Aucsia-silenced tomato plants exhibited auxin-related phenotypes such as parthenocarpic fruit development, leaf fusions, and reflexed leaves. Auxin-induced rhizogenesis in cotyledon explants and polar auxin transport in roots were reduced in Aucsia-silenced plants compared with wild-type plants. In addition, Aucsia-silenced plants showed an increased sensitivity to 1-naphthylphthalamic acid, an inhibitor of polar auxin transport. We further prove that total indole-3-acetic acid content was increased in preanthesis Aucsia-silenced flower buds. Thus, the data presented demonstrate that Aucsia genes encode a novel family of plant peptides that control fruit initiation and affect other auxin-related biological processes in tomato. Aucsia homologous genes are present in both chlorophytes and streptophytes, and the encoded peptides are distinguished by a 16-amino-acid-long (PYSGXSTLALVARXSA) AUCSIA motif, a lysine-rich carboxyl-terminal region, and a conserved tyrosine-based endocytic sorting motif.     

Effect of prolonged root hypoxia on the antioxidant content of tomato fruit

Tomato fruit quality depends on its metabolite content, which in turn is determined by numerous metabolic changes occurring during fruit development and ripening. The aim of this work was to investigate whether flooding affects the nutritional quality of tomato fruit, focusing on compounds essential to human health: carotenoids and ascorbate. To this end, tomato plants (Solanum lycopersicum L. cv Micro-Tom) were submitted to prolonged root hypoxia (1–2% O₂) at first flower anthesis. Fruits were harvested at five stages of the ripening process and analysed for their carotenoid and ascorbate contents. Our results showed that the ripening of fruits that developed on hypoxia treated plants was not inhibited. However, root hypoxia significantly limits carotenoid and ascorbate accumulation in pericarp during fruit ripening, the strongest effects being observed at late stages of ripening. Limitation of both carotenoids and ascorbate accumulation seems to be primarily mediated by the reduced level of expression of genes of the corresponding metabolic pathway.     

疏花对富士苹果光合同化物分配的外部调节和生理表现

在盛花期,利用对树冠不同部位疏除花序的试验。对富士苹果树体内同化物的调配及其生理表现进行了研究。结果表明,苹果平均单果重和平均单果叶面积之间关系呈线性正相关。该相关关系在不同处理之间无差异（r=0.91),充分说明在苹果树体内存在一个可自由流动的碳水化合物库或者说树体对结果具有光合调节作用,疏花可以从外部调节光合同化物分配的方向。对每株苹果树未结果部位和结果部位短枝及延长枝叶片净光合速率测定表明。结果和未结果部位光合速率没有显著差异。虽然疏花没有显著影响苹果单株枝条生长量和叶面积,但是,苹果树树冠全部疏除花序的一侧 或主枝的平均枝条生长量和叶面积去比该树其它结果部位分别平均高出153.5cm和8900cm^2,差异极为显著。苹果树疏除花序1/2或3/4的数量使苹果平均单果重比不疏花分别显著增加了33g或79g。而果实钙浓度每百克鲜重分别降低了1.62mg或2.66mg,充分说明当疏花显著地影响果实大小时,果实内钙浓度也会明显地发生相应的变化,虽然在疏花处理之间不同部位疏除花序没有影响单株花芽总数量,但是,每株树前一年春季疏花的部位比未疏花,结果的部位极显著地增加了花芽数量（SED=11.6)。     

A model for an early stage of tomato fruit development: cell multiplication and cessation of the cell proliferative activity

Changes in cell number during the early period of tomato fruit development were analysed by means of a deterministic model of cell multiplication. The period commenced at the seed stage with one theoretical cell undergoing intensive cell division, and ended when the cell number became nearly constant. The model takes into consideration the proliferative activity of the fruit cell population which, a few days before flower anthesis, begins to decrease progressively after each mitotic cycle. Model parameters, namely the time at which proliferative activity diminishes, its rate of decrease and the length of the cell cycle, were estimated by fitting the model to observed cell population dynamics in tomato fruits growing in three different positions on the truss. It is hypothesized that the molecular mechanism responsible for the cessation of mitosis in growing fruits is associated with shortening of telomeric ends of nuclear DNA, as suggested previously for other growing cell populations.     

Silencing of DELLA induces facultative parthenocarpy in tomato fruits

DELLA proteins are plant nuclear factors that restrain growth and proliferation in response to hormonal signals. The effects of the manipulation of the DELLA pathway in the making of a berry-like fruit were investigated. The expression of the Arabidopsis thaliana gain-of-function DELLA allele Atgai (del) in tomato (Solanum lycopersicum L.) produced partially sterile dwarf plants and compacted influorescences, as expected for a constitutively activated growth repressor. In contrast, antisense silencing of the single endogenous tomato DELLA gene homologue (SlDELLA) produced slender-like plants with elongated flower trusses. Interestingly, the depletion of SlDELLA in tomato was sufficient to overcome the growth arrest normally imposed on the ovary at anthesis, resulting in parthenocarpic fruits in the absence of pollination. Antisense SlDELLA-engineered fruits were smaller in size and elongated in shape compared with wild type. Cell number estimations showed that fruit set, resulting from reduced SlDELLA expression, arose from activated cell elongation at the longitudinal and lateral axes of the fruit pericarp, bypassing phase-II (post-pollination) cell divisions. Parthenocarpy caused by SlDELLA depletion is facultative, as hand pollination restored wild-type fruit phenotype. This indicates that fertilization-associated SlDELLA-independent signals are operational in ovary-fruit transitions. SlDELLA was also found to restrain growth in other reproductive structures, affecting style elongation, stylar hair primordial growth and stigma development.     

Dry Matter Partitioning in Tomato: Validation of a Dynamic Simulation Model

A model for dynamic simulation of dry matter distribution betweenreproductive and vegetative plant parts and the distributionamong individual fruit trusses in glasshouse tomato, is validated.The model is part of the crop growth model TOMSIM and is basedon the hypothesis that dry matter distribution is regulatedby the sink strengths of the plant organs, quantified by theirpotential growth rates, i.e. the growth rates at non-limitingassimilate supply. Within the plant, individual fruit trussesare distinguished and sink strength of a truss is describedas a function of its development stage. Truss development rateis a function of temperature only. The same potential growthcurve, proportional to the number of fruits per truss, is adoptedfor all trusses. In a simple version of the model, vegetativeplant parts are lumped together as one sink with a constantsink strength. In a more detailed version, vegetative sink strengthis calculated as the sum of sink strengths of vegetative units(three leaves and stem internodes between two trusses). The model was validated for six glasshouse experiments, coveringeffects of planting date, plant density, number of fruits pertruss (pruning at anthesis), truss removal (every second trussremoved at anthesis), single- and double-shoot plants and atemperature experiment conducted in climate rooms at 17, 20or 23 °C. Daily increase in above-ground dry weight, averagedaily temperatures and number of set fruits per truss were inputsto the model. Both the simple and the more detailed model showedgood agreement between measured and simulated fraction of drymatter partitioned into the fruits over time. For the simpleversion of the model, the slope of the lines relating simulatedto measured fraction partitioned into the fruits (16 data sets),varied between 0.92 and 1.11, on average it was 1.04, implying4% over-estimation for this fraction. For the detailed modelthese numbers were slightly better: 0.89, 1.08 and 1.01, respectively.The temperature experiment revealed no important direct influenceof temperature on the ratio between generative and vegetativesink strength. Simulated truss growth curves showed reasonableagreement with the measurements, although both models over-estimated(17% on average) final dry weight of the lower trusses (truss1 –3) on a plant. Modelling dry matter partitioning basedon sink strengths of organs is promising, as it is a general,dynamic and flexible approach, showing good agreement betweenmeasurements and simulation for a range of conditions. Applicabilityof the model is, however, still limited as long as the numberof fruits per truss (flower and /or fruit abortion) is not simulated,as this is a major feedback mechanism in plant growth. Dry matter distribution; sink strength; glasshouse; model; partitioning; simulation; temperature; tomato; TOMSIM; validation     

Gibberellins Increase Cuticle Deposition in Developing Tomato Fruit

Effects of the gibberellins A₄₊₇(GA₄₊₇) and A₃(GA₃), benzyladenine (BA) and forchlorfenuron (CPPU) on deposition of the cuticular membrane (CM) in developing tomato (Lycopersicon esculentum L.) fruit were investigated. Growth regulators were applied when fruit development within trusses ranged from the flower to the mature stage. Developmental stage of fruit at the time of application was indexed by fruit diameter. Fruit were harvested at maturity, the CM isolated enzymatically on an individual fruit basis and mass of CM per unit fruit surface area calculated. In mature fruit, mass of CM per fruit increased with fruit size, but mass of CM per unit surface area was independent of fruit size, position within a truss and position of the truss on the plant. GA₄₊₇ and GA₃ increased CM mass per unit fruit surface area at concentrations up to 300 mg l⁻¹. Young fruit (5–10 mm diam. at time of application) was most responsive. Responsiveness decreased as fruit development at application progressed towards maturity. There was no consistent effect of GA₄₊₇ or GA₃ on fruit mass. BA (up to 100 mg l⁻¹) or CPPU (up to 3 mg l⁻¹) had no significant effect on CM mass per unit surface area regardless of developmental stage. Higher concentrations of BA or CPPU decreased CM mass per unit surface area. There was no effect of BA or CPPU on fruit mass. Potential mechanisms and benefits of a gibberellin induced increase in CM deposition are discussed.     

Influence of Sink-Source Interaction on Dry Matter Production in Tomato

Sink-source ratio in tomato was manipulated, in six glasshouseexperiments, by fruit pruning (trusses pruned to two to sevenfruits immediately after fruit set of each truss), truss pruning(removal of every other truss at anthesis) and truss pruningin plants with two shoots. Periodic destructive harvest wereconducted for about 100 d after flowering of the first truss.Dry matter production was not influenced by sink-source ratio,whereas dry matter distribution between fruits and vegetativeparts was greatly affected. The fraction of dry matter distributedto the fruits at the end of the fruit pruning experiments (F_fruits)could be described accurately as a saturation-type functionof number of fruits retained per truss (N_f): F_fruits = 0.660(l-e^-0.341N_f). Specific leaf area and internode length decreasedand plant leaf area increased when sink-source ratio was reduced.Removal of every other truss at anthesis did reduce dry matterpartitioning into the fruits, but it did not influence internodelength. Plant development (number of visible leaves at the endof the experiments) was not influenced by sink-source ratio.In four experiments some plants were pruned to one fruit pertruss. Final dry matter production was 8-24% lower for theseplants, compared with plants with more than one fruit per truss.This was, at least party, the result of less light interceptionby these plants, which had strongly curled leaves pointing downwards. Results indicate that effects of sink demand on dry matter productionper unit of intercepted radiation and probably on leaf photosyntheticrate in commercial tomato production can be ignored.Copyright1995, 1999 Academic Press Dry matter production, feedback control, glasshouse, growth analysis, Lycopersicon esculentum, pruning, sink demand, sink-source ratio, tomato     

The specific overexpression of a cyclin-dependent kinase inhibitor in tomato fruit mesocarp cells uncouples endoreduplication and cell growth

The size of tomato fruit results from the combination of cell number and cell size, which are respectively determined by the cell division and cell expansion processes. As fruit growth is mainly sustained by cell expansion, the development of fleshy pericarp tissue is characterized by numerous rounds of endoreduplication inducing a spectacular increase in DNA ploidy and mean cell size. Although a clear relationship exists between endoreduplication and cell growth in plants, the exact role of endoreduplication has not been clearly elucidated. To decipher the molecular basis of endoreduplication-associated cell growth in fruit, we investigated the putative involvement of the tomato cyclin-dependent kinase inhibitor SlKRP1. We studied the kinetics of pericarp development in tomato fruit at the morphological and cytological levels, and demonstrated that endoreduplication is directly proportional to cell and fruit diameter. We established a mathematical model for tissue growth according to the number of divisions and endocycles. This model was tested in fruits where we managed to decrease the extent of endoreduplication by over-expressing SlKRP1 under the control of a fruit-specific promoter expressed during early development. Despite the fact that endoreduplication was affected, we could not observe any morphological, cytological or metabolic phenotypes, indicating that determination of cell and fruit size can be, at least conditionally, uncoupled from endoreduplication.