Relationship between cell number,cell size and fruit size of seeded fruits of tomato (Lycopersicon esculentum Mill.), and those induced parthenocarpically by the application of plant growth regulators

Application of GA₃, IAA or 4-CPA to tomato ovaries induced the development of parthenocarpic fruit, which showed different growth rates. In the pericarp cell division and cell enlargement was affected differentially. GA₃-induced fruits had considerably less but larger cells than seeded control fruits, IAA treatment resulted in the same number of cells but these were smaller and 4-CPA treatment induced fruits with about 20% more cells. Reduction in cell number had a similar effect on final fruit size as diminution of cell size. A reduction in the number of cell division centres (area around vascular bundles) as well as changes in the degree of endoploidy are possible reasons for the observed reductions in cell numbers. Hormonal causes for the different number and size of pericarp cells after the various treatments are discussed.     

Do Genetic Make-up and Growth Manipulation Affect Tomato Fruit Size by Cell Number,or Cell Size and DNA Endoreduplication?

This work investigated the link between genetic and developmental controls of fruit size and composition. On two isogenic lines (CF12-C and CF14-L), differing by fruit weight and sugar content quantitative trait loci (QTLs) identified previously, basal and tip fruits were characterized at anthesis and at maturity through their growth, dry matter and sugar content, number and size of cells and nuclei DNA content. The influence of competition was assessed by removing either basal or tip ovaries at anthesis. On an intact inflorescence, CF12-C fruits grew less than CF14-L fruits, with 1.67 fewer cell layers and similar cell size, suggesting that genes controlling cell division may be responsible for this fruit size variation. Truss thinning masked the QTL effect on fruit size, mainly by reducing the difference in cell number between the two lines and by promoting cell expansion in tip fruits, so that fruit growth was similar at both positions and for both lines. Thus, in these lines, cell number exerts a control on final fruit size only when there is competition among fruits. Different responses of basal and tip fruits after flower removal suggested that this treatment induced changes in hormonal relationships within the truss. No fixed relationship between DNA endoreduplication and cell size was found, as while cell size and dry matter and sugar contents differed with tomato lines, fruit position and truss size, endoreduplication patterns were the same. CF12-C fruits had a higher dry matter (+0.3% of fresh weight) and carbohydrates (+8% of dry matter) content than CF14-L fruits. The percentage dry matter was independent of truss size but decreased slightly from basal to tip fruits.     

Analysis of the tomato fruit growth response to temperature and plant fruit load in relation to cell division, cell expansion and DNA endoreduplication

BACKGROUND AND AIMS: To better understand the regulation of fruit growth in response to environmental factors, the effects of temperature and plant fruit load on cell number, cell size and DNA endoreduplication were analysed. METHODS: Plants were grown at 20/20 degrees C, 25/25 degrees C and 25/20 degrees C day/night temperatures, and inflorescences were pruned to two ('2F') or five ('5F') flowers. KEY RESULTS AND CONCLUSIONS: Despite a lower fruit growth rate at 20/20 degrees C, temperature did not affect final fruit size because of the compensation between cell number and size. The higher cell number at 20/20 degrees C (9.0 x 10(6) against 7.9 x 10(6) at 25/25 degrees C and 7.7 x 10(6) at 25/20 degrees C) resulted from an extended period of cell division, and the smaller cell size was due to a shorter period of expansion rather than a lower expansion rate. By contrast, the lower fruit growth rate and size of 5F fruits compared with 2F fruits resulted from the slow down of cell expansion, whereas the number of cells was hardly affected in the proximal fruit. However, within the inflorescence the decreasing gradient of fruit size from proximal to distal fruits was due to a decrease in cell number with similar cell size. Fruit size variations within each treatment were always positively correlated to variations in cell number, but not in cell size. Negative correlations between cell size and cell number suggested that cells of tomato pericarp can be seen as a population of competing sinks. Mean ploidy was slightly delayed and reduced in 5F fruits compared with 2F fruits. It was highest at 25/25 degrees C and lowest at 25/20 degrees C. Treatments did not affect ploidy and cell size in similar ways, but within each treatment, positive correlations existed between mean ploidy and cell size, though significant only in the 2F-25/20 treatment.     

Cell number, cell size and hormone levels in semi-isogenic mutants of Lycopersicon pimpinellifolium differing in fruit size

Tomato fruit growth parameters, cell number and cell size, and hormone levels [IAA, abscisic acid (ABA), zeatin (Z)/zeatin riboside (ZR), isopentenyladenosine (i-Ado)/isopentenlyadenine (i-Ade)], in the wild-type ( Lycopersicon pimpinellifolium Mill.) and a semi-isogenic mutant (mutant III) differing in fruit size were investigated during fruit development. An image-processing system was used for the determination of cell number and single cell size per fruit and hormone levels were measured by radioimmuno-assay (RIA). The bigger fruits of mutant III showed higher cell numbers throughout fruit development and cells enlarged faster than in wild-type fruits. During the first 10 days of fruit growth, the main cell division period after fertilization, high concentrations of cytokinins were found, these being correlated with high cell division activity. There were only slight differences in IAA and ABA levels in the different sized fruits. The results emphasized the importance of the cell number per fruit at anthesis as a determining factor of final fruit size in tomatoes. A possible relationship between cytokinins and subsequent fruit development is discussed.     

Increased activity of ornithine decarboxylase in tomato ovaries induced by auxin

Auxin, which mimics natural pollination in tomato flowers, induces fruit setting as well as a 3-fold increase in ornithine decarboxylase activity. In pollinated ovaries ornithine decarboxylase activity increased 4.7 fold. The possible connection between pollination, auxin treatment, fruit set, cell division and ornithine decarboxylase activity are discussed.     

Growth and development of tomato fruitsin vivo andin vitro

Tomato (Lycopersicon esculentum Mill.) fruits of the male sterile cultivar Pearson (MS35 BC4, 61) were transferred toin vitro culture during the cell division period. Fruits grownin vivo andin vitro were compared throughout their development according to various growth parameters: fresh and dry weight, cell number, cell diameter, and DNA and total protein content. In all cases, the values pertaining to fruits grownin vitro were significantly lower than theirin vivo counterparts. The final fresh weight of fruits transferred to culture 2, 5, or 10 days after pollination was only 0.7, 1.2, and 3.4%, respectively, of that of plant-grown fruits. The results indicate that the reduced fruit sizein vitro is related to the reduction in both cell number and cell size. It is interesting to note that the DNA content per cell increased 15-fold during the growth of the plant-grown fruits while this accumulation was only between 2-and 3-fold in all the cultured fruits. The time to first colour appearance of fruits cultured 2, 5, or 10 days after pollination was 196, 132 and 85%, respectively, of that of plant-grown fruits.     

Effect of Gibberellin and Auxin on Parthenocarpic Fruit Growth Induction in the cv Micro-Tom of Tomato

The effect of applied gibberellin (GA) and auxin on fruit-set and growth has been investigated in tomato (Solanum lycopersicum L.) cv Micro-Tom. It was found that to prevent competition between developing fruits only one fruit per truss should be left on the plant. Unpollinated ovaries responded to GA₃ and to different auxins [indol-3-acetic acid, naphthaleneacetic acid, and 2,4-dichlorophenoxyacetic acid (2,4-D)], 2,4-D being the most efficient. GA₃- and 2,4-D-induced fruits had different internal morphology, with poor locular tissue development in the case of GA, and pseudoembryos development in the case of 2,4-D. Also, GA₃ produced larger cells in the internal region of the mesocarp (IM) associated with higher mean C values, whereas 2,4-D produced more cell layers in the pericarp than pollinated fruits. The smaller size of GA₃- compared with 2,4-D-induced fruits was due to them having fewer cells, only partially compensated by the larger size of IM cells. Simultaneous application of GA₃ and 2,4-D produced parthenocarpic fruits similar to pollinated fruits, but for the absence of seeds, suggesting that both kinds of hormones are involved in the induction of fruit development upon pollination. It is concluded that Micro-Tom constitutes a convenient model system, compared to tall cultivars, to investigate the hormonal regulation of fruit development in tomato.     

Blossom-end rot in relation to growth rate and calcium content in fruits of sweet pepper (Capsicum annuum L.)

The relative importance of growth rate and calcium concentration in sweet pepper fruits (Capsicum annuum L.) for the induction of blossom-end rot (BER) was investigated in (1) four pollination treatments in one cultivar, (2) four cultivars with the same fruit load and (3) three fruit load treatments in four cultivars. For fruits with the same pollination treatment those eventually developing BER had a higher initial fruit growth rate than those not developing BER. Within the same experiment both the growth rate of the young fruit and BER increased with the number of seeds. The Ca concentration of the pericarp in mature fruits was negatively related to both fruit size and BER incidence. Differences in levels of BER between different pollination experiments could not be explained solely by differences in growth rate of the young fruit, but related to different Ca concentrations in the mature fruits. In the spring, but not in the summer, cultivars more susceptible to BER had a larger final size but lower Ca concentration in the young fruit than the resistant ones. By lowering the fruit load in the summer both the final fruit size and the BER incidence increased, but the Ca concentrations of both proximal and distal pericarp in the young fruit of all cultivars were not consistently affected. Despite a correlation between growth rate and low Ca concentration in the fruit, the incidence of BER may only be predicted from separate effects of fruit growth and of Ca concentration of fruit. The data indicated that at a higher growth rate a higher Ca concentration is required to prevent the induction of BER. The usefulness of the total Ca concentration of the fruit for determining the critical Ca concentration in the induction of BER is discussed.Key words: Capiscum annuum L., sweet pepper, blossom-end rot, calcium, growth rate, pollination, fruit load.      

Polyamine levels in pollinated and auxin-induced fruit of tomato (Lycopersicon esculentum) during development

The changes taking place during fruit development in the concentration of the 3 polyamine fractions, i.e. free, perchloric acid-soluble conjugates and perchloric acid-insoluble bound polyamines, were analyzed in tomato fruits ( Lycopersicon esculentum Mill, cv. F121) induced to set by either pollination or auxin application. Before the onset of cell division, total polyamines were 50% higher in auxin-treated fruits than in pollinated ones, most of the polyamines being found as perchloric acid-soluble conjugates in both fruit set treatments. At the onset the level of polyamines in both fruit types was 100 times higher than during cell expansion or ripening. The highest polyamine found during cell division was perchloric acid-soluble conjugated spermidine in both fruit set treatments. After cell division, polyamine concentration was similar in both fruit set treatments. The concentration of polyamines in the jelly was similar in pollinated and auxin-induced fruits during cell expansion but not during ripening. At the onset of ripening there was an increase of one order of magnitude in the concentration of perchloric acid-insoluble bound putrescine in the jelly of pollinated fruits. Polyamines were more than 5-fold higher in unpollinated ovaries than in fruits induced to set by either pollination or auxins. It is suggested that pollinated and parthenocarpic fruits differ in their polyamine metabolism during the initial stages of development, but not after cell division. It is also suggested that polyamines undergo rapid turnover during cell division. Perchloric acid-insoluble bound putrescine might play a role in seed formation in tomatoes.     

A rapid increase in cytokinin levels and enhanced ethylene evolution precede Al3+-induced inhibition of root growth in bean seedlings (Phaseolus vulgaris L.)

Fruit sink strength or its ability to attract assimilates depends bothon sink activity and size. This study investigated one main component of sinksize, that is the number of fruit cells during tomato plant development. Plantswere grown in a controlled climate chamber under a limiting (LS, six fruits pertruss) and non-limiting (NLS, two fruits per truss and CO₂enrichment) supply of carbon assimilates. Under NLS conditions, fruit cellnumber was homogeneous among successive trusses, and fruits contained onaverage1.2×10⁶ more cells than under LS conditions,though differences were not significant on the first truss which underwent thelowest competition. Under LS conditions, an ontogenetic increase in cell numberwas observed in proximal fruits of the upper trusses attributed to theenlargement of the apical meristem during plant development. The decrease ofcell number from proximal to distal fruits within a truss, that was expectedfrom the literature, was generally observed in the LS experiment, with anaverage significant difference of about1.6×10⁶cells between the first and fifth fruits. Nevertheless, whereas the gradient incell number from proximal to distal fruits was steep in the upper trusses, itwas not significant on the lower trusses indicating that this gradient largelydepended on the level of competition during floral development. Thus, under lowassimilate supply, cell division is a main limiting factor for fruit growth,although cell enlargement during further fruit development is also affected,butwas not measured in this work.     

The effect of difluoromethylornithine on polyamine levels in pollinated and napthaleneacetic acid-induced young tomato fruits

The polyamine synthesis inhibitor difluoromethylornithine (DFMO) inhibits growth and cell division in tomato fruits (Lycopersicon esculentum Mill. c.v. F 121) when applied after pollination by the split-stem technique. In napthaleneacetic acid (NAA)-induced fruits, however, growth is not inhibited by DFMO. We analyzed the effect of DFMO on cell division in auxin induced tomato fruits, and on the concentration of free, soluble conjugated and insoluble bound polyamines in pollinated and NAA-induced fruits, five days after pollination in the presence and absence of DFMO. Cell number was not significantly reduced by DFMO in NAA-induced fruits five days after fruit set. Free spermine, soluble conjugated and insoluble bound polyamines were higher in fruits treated with DFMO than in those treated with water. They were also higher in DFMO-treated pollinated fruits than in NAA-induced ones. Our results suggest that inhibition of cell division by polyamine synthesis inhibitor might affect further metabolism of polyamines.Abbreviations DFMA -difluoromethylarginine - DFMO -difluoromethylornithine - MGBG methylglyoxyl bis-(guanylhydrazone) - NAA -naphthaleneacetic acid Department of Life Sciences Ben-Gurion University of the NegevThis work was performed in partial fulfillment for the PhD degree of Marcos Egea-Cortines     

大花蕙兰与墨兰杂交结实率研究初报

取3个大花蕙兰品种与1个墨兰品种进行自交与杂交。共设计10个组合,成功获得4个杂交组合;共授粉257朵花,成功获得21个果荚,结实率为8.1%。大花蕙兰与墨兰杂交正反交的结实率差异不大。各组合果荚生长曲线走势大致相同,授粉后10~12 d,子房开始膨大,并于60~80 d趋于平稳。大部分落果于授粉后30d内发生,部分于生长中后期黄化脱落,双亲遗传物质的不协调性及栽培管理可能是重要影响因素。     

Enhancement of fruit quality in Capsicum annum through pollination by Hypotrigona gribodoi in Kakamega,Western Kenya

A study was carried out in Kakamega forest, in the western region of Kenya, to evaluate the effectiveness of the stingless bee Hypotrigona gribodoi (Magretti, 1884) on the pollination of green pepper. Three treatments were applied and consisted of self‐pollination, pollination by feral pollinators in the open field and pollination by H. gribodoi in a net cage. The differences in fruit yield and seed quality were compared among treatments. Flowers pollinated by H. gribodoi produced the heaviest fruits with the highest seed numbers followed by feral pollinators and lastly self‐pollinated flowers. Moreover, seeds were significantly bigger in size in fruits resulting from flowers pollinated by H. gribodoi compared to fruits obtained from self‐pollinated flowers or flowers pollinated by feral insects. We, therefore, conclude that H. gribodoi is an efficient pollinator of green pepper in the tropical region of East Africa.     

Fruit size decline from the margin to the center of capitula is the result of resource competition and architectural constraints

Plants produce repeated structures, such as leaves, flowers, and fruits, which differ in size and shape. One example of this is fruit size, which is commonly observed to decrease from proximal to distal positions within an inflorescence. The resource limitation hypothesis proposes that because proximal fruits usually develop first, they have temporal priority on access to resources over distal fruits. The non-mutually exclusive architectural effects hypothesis suggests that these position effects in fruit size may also be due to inherent architectural variation along infructescence axes. We separated out the effects of resource competition and inflorescence architecture by removing the outer or the inner flowers within capitula of Tragopogon porrifolius. We also studied if fruit position influenced germination and seedling performance in order to assess fitness consequences of position effects. Inner fruits were significantly heavier when outer flowers were removed. However, outer fruits did not significantly increase when inner flowers were removed, suggesting later fruits were limited by the development of early fruits. Our findings also suggest that architectural constraints restricted the size of inner fruits in comparison with outer ones. We found that both resource competition and inflorescence architecture affected the fruit size of T. porrifolius, even though this species does not have linear, indeterminate inflorescences. We advance the hypothesis that, when such effects on fitness occur, resource competition-mediated position effects could turn, in evolutionary time, into architectural position effects.     

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.     

Early Application of Ethrel Extends Tomato Fruit Cell Division and Increases Fruit Size and Yield with Ripening Delay

Flowers of tomato (Lycopersicon esculentum Mill.) plants cv. Castle Rock were sprayed with 100 ppm of ethrel, 0.5 mm aminooxyacetic acid (AOA), or water (control) 2 days after anthesis. The fruit period of cell division was extended up to 16–18 days after anthesis with the application of ethrel but reduced from 10–12 days (control) down to only 6–8 days with the application of AOA. In a trend opposite to AOA application, fruits that received ethrel treatment were of higher ethylene and 1-aminocyclopropane-1-carboxylic acid (ACC) levels than control. This was noticed not only during the first 2 weeks after anthesis but also during the fruit climacteric phase. Mesocarp cells of ethrel-treated fruits were greater in number/mm² but smaller in size than control; an opposite trend was obtained with the application of AOA. This was observed for a period of 18 days after anthesis, but by that time or at earlier ages, fruits of AOA treatment were larger in size and heavier in weight than control, and both were larger and heavier than ethrel-treated ones. At 5 weeks after anthesis and thereafter, the fruit response to all treatments was totally reversed because early ethrel-treated fruits became significantly larger in size and heavier in weight with a ripening delay of about 10 and 15 days compared with those of control and AOA-treated ones, respectively. When the same treatments were applied to the whole plant, similar results were obtained because the early application of ethrel increased the fruit yield by about 15% over control with a pronounced ripening delay; an opposite trend was obtained with the application of AOA. No significant differences were found among all treatments in terms of flower or fruit abscission or fruit number/plant. The data suggest that ethylene regulates tomato fruit transmission from cell division to cell enlargement. In addition, fruit cell division is terminated only when endogenous ethylene decreases to its basal level, allowing cell enlargement to dominate and proceed as in the case of the early application of AOA. The ripening delay of ethrel-treated fruits may be caused by the longer time required for the increased cell number to reach maturation. A low level of ethrel application at the tomato early fruiting stage may be used for increasing fruit yield by increasing fruit size and consequently its quality. Received June 1, 1998; accepted December 7, 1998     

Cell division and cell enlargement in fruit of Lagenaria leucantha as influenced by pollination and plant growth substances

The effects of NAA (naphthaleneacetic acid), GA₃ (gibberellic acid), CPPU (N-(2-chloro-4-pyridyl)-N'-phenylurea) and pollination on fruit set, cell division and enlargement were studied in Lagenaria leucantha, an important vegetable. NAA and GA₃ were ineffective in inducing parthenocarpy, whereas CPPU induced parthenocarpic fruit significantly larger than fruit that resulted from pollination. Cell division, which occurred during the first 4 days after pollination was not reactivated by NAA or GA₃, but was effectively reactivated by CPPU. The cell number of the total cross-section of CPPU-treated fruit was 117.4% of that of pollinated fruit and 154.4% of that of unpollinated at 12 DAA (days after anthesis) respectively. The CPPU-induced parthenocarpic fruit had the largest cell cross-sectional area followed, successively, by pollinated fruit, NAA-treated fruit, GA₃-treated fruit and unpollinated fruit. These results indicate that CPPU induced parthenocarpic fruit growth by directly reactivating cell division and expansion.     

The impact of cell division and cell enlargement on the evolution of fruit size in Pyrus pyrifolia

BACKGROUND AND AIMS: Dramatic increases in fruit size have accompanied the domestication of Pyrus pyrifolia. To evaluate the contribution of cell division and cell enlargement in the evolution of fruit size, the following study was conducted. METHODS: Three wild Pyrus and 46 cultivated Pyrus pyrifolia cultivars were selected to examine cell number/size at time of pollination and at time of fruit harvest. The period of cell division was estimated by logarithmic curve of the increasing pattern of cell number, and its correlations with maturation period and final fruit size were analysed. KEY RESULTS: Final fruit size is directly related to the number of cells produced in the period immediately following pollination. Late-maturing cultivars are larger than earlier-maturing cultivars and this is due to an extended period of cell division. CONCLUSIONS: The evolution of fruit size in P. pyrifolia has mainly resulted from shifts in the ability of cells to divide rather than to enlarge.     

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.