Fruit Position and Fruit Set Sequence in a Truss as Factors Determining Final Size of Tomato Fruits

The position of fruits in a truss and the induction sequenceof fruit set has been examined for the effects on the finalsize of parthenocarpic fruits in tomato. Fruits at two developmentalstages were examined to measure the change of d. wt and theimport of ¹⁴C leaf assimilates. The sequence of induction largely determined the import rateinto fruits of the same truss and first-induced fruits are thereforelarger within the same truss. The position of fruits in thetruss modifies the extent of this ‘primigen dominance’effect. Possible mechanisms for this dominance phenomenon arediscussed. Position, set sequence, size, fruit, Lycopersicon esculentum Mill     

Relationship between set, development and activities of growth regulators in tomato fruits

The actions of plant regulators in set and development of fruitsare well known. However, the presence and function of endogenoushormones in parthenocarpic fruits have still not been sufficientlyinvestigated. A comparison between seeded and seedless fruitsmakes it possible to obtain a more accurate understanding ofsome relationships between growth regulators and stages of fruitdevelopment. Endogenous auxin and gibberellin activity levelsand some growth parameters (fresh and dry weight, cell numberand cell volume, DNA content) have been determined in tomatofruits (Lycopersicon esculentum Mill.) of the cultivar Venturaand of its isogenic parthenocarpic mutant. In both genotypes,auxin and gibberellin are present in the first week after anthesis,though at different concentrations and with different patterns.These two activities are involved in fruit setting. The simultaneousoccurrence of maximum auxin concentration and of the beginningof cell enlargement, in both genotypes, shows that the activitypresent at this time starts fruit development and possibly determinesthe size of the fruits. High auxin activity is observed only in seeded fruits 20–40days after anthesis, and it is probably synthesized by seeds.Gibberellin activity is present, corresponding to the changein fruit development from the mature green to pink stages. (Received February 16, 1978; )     

Induced parthenocarpy-a way of manipulating levels of endogenous hormones in tomato fruits (Lycopersicon esculentum Mill.) 2. Diffusible hormones

Indol-3-acetic acid (IAA), gibberellin-like substances (GAs), and abscisic acid (ABA) were measured throughout the first 35 days of fruit development in agar diffusates from seeded and parthenocarpic tomato fruits. Parthenocarpic fruit growth was induced with either an auxin (4-CPA), morphactin (CME) or gibberellic acid (GA₃). IAA and GAs were at their highest levels in diffusates during the early stages of fruit growth, whereas diffusible ABA increased later. Most IAA was found in diffusates from auxin-induced and seeded fruits, whereas GAs were at their lowest levels in seeded fruits. There were only minor differences in ABA concentrations regardlesss of the treatments.Levels of diffusible hormones of tomato fruits may be easily manipulated by inducing parthenocarpic fruit growth. In spite of no obvious relationship between fruit growth and hormone levels in this study, induced parthenocarpy is considered a useful tool to further elucidate the role of hormones in fruit development and sink-source interactions.     

Parthenocarpy and seed predation by insects in Bursera morelensis

Background and Aims

While parthenocarpy (meaning the production of fruits without seeds) may limit fecundity in many plants, its function is not clear; it has been proposed, however, that it might be associated with a strategy to avoid seed predation. Bursera morelensis is a dioecious endemic plant that produces fruits with and without seeds, and its fruits are parasitized by insects. Its reproductive system is not well described and no published evidence of parthenocarpy exists for the species. The purpose of this work was to describe the breeding system of B. morelensis and its relationship to seed predation by insects.

Methods

The breeding system was described using pollination experiments, verifying the presence of parthenocarpic fruits and apomictic seeds. Reproductive structures from flower buds to mature fruits were quantified. For fruits, an anatomical and histological characterization was made. The number of fruits in which seeds had been predated by insects was correlated with parthenocarpic fruit production.

Key Results

The major abortion of reproductive structures occurred during fruit set. The results discard the formation of apomictic seeds. Flowers that were not pollinated formed parthenocarpic fruits and these could be distinguished during early developmental stages. In parthenocarpic fruits in the first stages of development, an unusual spread of internal walls of the ovary occurred invading the locule and preventing ovule development. Unlike fruits with seeds, parthenocarpic fruits do not have calcium oxalate crystals in the ovary wall. Both fruit types can be separated in the field at fruit maturity by the presence of dehiscence, complete in seeded and partial in parthenocarpic fruits. Trees with more parthenocarpic fruits had more parasitized fruits.

Conclusions

This is the first time the anatomy of parthenocarpic fruits in Burseraceae has been described. Parthenocarpic fruits in B. morelensis might function as a deceit strategy for insect seed predators as they are unprotected both chemically and mechanically by the absence of calcium oxalate crystals.Key words: Parthenocarpy, Bursera morelensis, predation, seeds, insects, breeding system, calcium oxalate crystals     

Differential regulation of a fruit-specific 62 kDa protein in developing parthenocarpic (pat-2/pat-2) and seeded tomato fruits

The denatured protein profiles of developing tomato ( Lycopersicon esculentum Mill.) fruits, from the anthesis stage up to fruits at 30% of their final diameter, were examined in a pai-2l pat-2 parthenocarpic line and in its near isogenic non-partheno-carpic line. At anthesis no differences were detected between the protein patterns of ovaries developed on parthenocarpic and non-parthenocarpic plants. In subsequent stages the seeded and seedless fruits differed in the pattern of manifestation of several abundant proteins, none of which seem to be included in seeds The most prominent difference was found in an insoluble protein of 62 kDa; in developing seeded fruits of either the parthenocarpic or the non-parthenocarpic line, its rate of decline was much faster than in seedless fruits. In seeded fruits larger than 4-6 mm in diameter it was scarcely detected, whereas in parthenocarpic seedless 8–10 mm fruits it was still abundant. This protein is fruit specific; it is also enhanced in chemically (n-n-tolyl phthalamic acid) – induced parthenocarpic fruits of the non-parthenocarpic line. The prolonged manifestation in the parthenocarpic fruits results from de novo synthesis of this protein. There are indications that it is not a stress-related protein. This is the first demonstration of an association between the pattern of modulation of a protein and the phenotypic expression of genetically controlled parthenocarpy.     

Induction of Parthenocarpy in Rosa Canina and Diospyros Lotus by the application of growth regulators

The effect of indole-3-acetic acid (IAA) and gibberellic acid (GA₃) applications on parthenocarpic fruit set in Rosa canina and Diospyros lotus was investigated. GA₃ induced parthenocarpic fruit set in both plants, but IAA only in D. lotus. Maturation of seedless fruits was earlier than the seeded fruits. GA₃ caused a decrease in the fresh mass and size of both fruits. IAA induced an increase in the fresh mass and size in parthenocarpic fruit of D. lotus. This revised version was published online in July 2006 with corrections to the Cover Date.     

Induced parthenocarpy—a way of changing the levels of endogenous hormones in tomato fruits (Lycopersicon esculentum Mill.) 1. Extractable hormones

Ethylene, indol-3-acetic acid (IAA), gibberellin-like substances (GAs) and abscisic acid (ABA) were analysed in extracts from normal, seed-containing and parthenocarpic tomato fruits throughout fruit development. Parthenocarpic fruit growth was induced with an auxin (4-CPA), morphactin (CME) or gibberellic acid (GA₃) and compared with that of pollinated control fruits. Fruit growth was only affected by the treatment with GA₃, decreasing size and fresh weight by 60%. The peak sequence of hormones during fruit development was ethylene-GAs-IAA-ABA. Seeded fruits contained the highest levels of IAA and ABA but the lowest levels of GAs. Also, in seeded fruits, a high proportion of IAA and ABA was found in the seeds whereas this was not the case for GAs.Hormone levels of tomato fruits may be successfully, easily and reproducibly altered by inducing parthenocarpic fruit growth and thus eliminating development of seeds which are a major source of hormone synthesis. In spite of markedly changed hormone levels, there was no obvious relationship between fruit growth and extractable hormones per se. However, the results indicate that a high ratio of GAs: auxins is unfavourable for growth of tomato fruits.     

植物激素对果树单性结实调控的研究进展

尽管种子与果实的发育过程密切相关,但由于无籽果实提高了果实的品质并延长了货架期,因此无籽果实受到生产者和消费者的广泛青睐。果实的生长受激素调控,而相关研究也初步揭示了激素在单性结实果实形成过程中的作用。本文对单性结实的概念、诱导单性结实的激素种类及功能、诱导机理,以及转基因技术在单性结实果实生产中的应用概况进行了综述,以期为果树单性结实的研究提供参考。     

Chlorophenoxyacetic acid and chloropyridylphenylurea accelerate translocation of photoassimilates to parthenocarpic and seeded fruits of muskmelon (Cucumis melo)

We compared the effect of p-chlorophenoxyacetic acid (p-CPA) and 1-(2-chloro-4-pyridyl)-3-phenylurea (CPPU) on parthenocarpic and seeded muskmelon (Cucumis melo) fruits in regards to fruit development and the transport of photoassimilates from leaves exposed to ¹⁴CO₂ to the developing fruits. Ten days after anthesis (DAA), the fresh weight, total ¹⁴C-radioactivity and contents of ¹⁴C-sucrose and ¹⁴C-fructose were higher in the CPPU-induced parthenocarpic fruits than in seeded fruits. However, at 35 DAA, fresh weight and sucrose content in mesocarp, placenta and empty seeds of the parthenocarpic fruits were lower than in seeded fruits. Also, total ¹⁴C-radioactivity and ¹⁴C-sugar content of the parthenocarpic fruits were lower as well as the translocation rate of ¹⁴C-photoassimilates into these fruits. Application of p-CPA to the parthenocarpic fruits at 10 and 25 DAA increased fresh weight and sugar content. Moreover, these treatments elevated the total ¹⁴C-radioactivity, ¹⁴C-sucrose content and the translocation rate of ¹⁴C-photoassimilates. The ¹⁴C-radioactivity along the translocation pathway from leaf to petiole, stem, lateral shoot and peduncle showed a declining pattern but dramatically increased again in the fruits. These results suggest that the fruit's sink strength was regulated by the seed and enhanced by the application of p-CPA.     

Seedlessness and Parthenocarpy inPistacia veraL. (Anacardiaceae): Temporal Changes in Patterns of Vascular Transport to Ovules

Pistacia vera‘Kerman’ (pistachio nut) typicallyproduces high numbers of seedless or blank fruits. Patternsof vascular transport into fruits and ovules were studied over3 years by following the movement of disodium fluorescein solutionfrom cut branches into developing fruitlets. Early in the season,vascular conductivity is intact through to the chalazal endof the ovule. Soon afterwards, the percentage of ovules withvascular conductivity through to the chalaza declines, and ina variable fraction of fruits, movement of the fluorochromesolution becomes blocked either at the placenta or in the funiculus.Six to 9 weeks after anthesis there is blockage in 90 (1 year)to 100% (2 years) of fruits. Subsequently, vascular conductivityresumes in 83.3% (3 year mean) of ovules, a percentage thatcorrelates well with the mean percentage of seeded nuts at harvest(77.5%). Ovules from fruits with dysfunctional vascular conductionearly in the season are smaller than those with fully functionalvascular tissue. At the time conductivity declines, a high percentageof those ovules with blocked vascular movement lack endospermand appear to be unfertilized; none of the ovules that retainfull vascular flow lack endosperm. Pollination using gamma-irradiatedpollen (⁶⁰Co, 1.0 kGy) led to a nearly three-fold increase inthe production of blank nuts. The results indicate that fluoresceintransport may be a valuable tool to predict the fate of ovules,and are consistent with the hypothesis that parthenocarpic fruitset may be an important factor in blank nut production in pistachio.Copyright1999 Annals of Botany Company Pistachio,Pistacia veraL., fluorescein, seed set, seedlessness, parthenocarpy, blanking, ovule, funiculus, chalaza, embryo.     

Induction of parthenocarpy in tomato via specific expression of the<Emphasis Type="Italic"> rolB</Emphasis> gene in the ovary

The molecular signals for the development of the ovary into fruit following ovule fertilization are not clear. However, in many species, including tomato ( Lycopersicon esculentum Mill.), auxins and auxin transport inhibitors can substitute for fertilization as activators of fruit set, suggesting that this plant hormone plays a key role in this process. In agreement, transgenes for auxin biosynthesis expressed under ovary- or ovule-specific promoters were shown earlier to enable parthenocarpic (i.e. seedless) fruit development. In the present study, we tested an alternative approach for the induction of parthenocarpy that is based on ovary-specific expression of the Agrobacterium rhizogenes-derived gene rolB. This gene was chosen because rolB transgenic plants manifest several syndromes characteristic of auxin treatment. Tomato plants transformed with a chimeric construct containing the rolB gene fused to the ovary- and young-fruit-specific promoter TPRP-F1 developed parthenocarpic fruits. Fruit size and morphology, including jelly fill in the locules of the seedless fruits, were comparable to those of seeded fruits of the parental line. Although it is not known whether ROLB signals for the same cassette of genes involved in fertilization-dependent fruit development, it clearly activates a battery of genes that enable successful completion of seedless fruit development in tomato.     

l-Ascorbic acid metabolism in parthenocarpic and seeded cherry tomatoes

The auxin treatment in tomato plants during anthesis has been extensively used for setting fruits in adverse climatic conditions (e.g., low temperatures and inadequate light), which is well known that reduces pollen availability and fertility. Since auxin application may affect fruit composition and quality, we examined l-ascorbic acid metabolism in seeded fruit (set by natural pollination) and parthenocarpic fruit (set by auxin) in cherry tomato cv. Conchita. Specifically, we studied the oxidized and total ascorbic acid contents, the expression of all characterized genes of l-ascorbic acid metabolism, the activity of ascorbate peroxidase and dehydroascorbate reductase and the immunolocalization of ascorbate peroxidase. Differences were detected between seeded and parthenocarpic fruits, in the expression of some of the genes of ascorbic acid metabolism. However, strong presence of l-ascorbic acid peroxidase protein was detected on the developing seeds. Our data indicate that induced parthenocarpy in auxin treated plants has a significant influence in ascorbic acid metabolism comparing to seeded tomato fruits.     

Peroxidase and IAA oxidase activities and peroxidase isoenzymes in the pericarp of seeded and seedless “Redhaven” peach fruit

Parthenocarpic peach fruit (Prunus persica L. Batsch., cv. Redhaven) were induced with 1-(3-chlorophthalimide)-cyclohexane carboxamide (AC 94377). The activities of soluble, and ionically and covalently bound peroxidase and indole-3-acetic acid (IAA) oxidase in the pericarp of both seeded and parthenocarpic fruit were determined from 21–43 days after anthesis. Seedless fruit grew faster during early stage I and ceased growth earlier than seeded fruit. Total peroxidase and IAA oxidase activities increased with development on both types of fruit, but higher values were found in seedless fruit. The ionic fraction showed the greatest increase for both enzyme activities. Isoperoxidase profile showed new cationic isoenzymes and higher levels of the less anionic isoenzymes in the pericarp of seedless fruit, whereas the seeded fruit contained higher levels of the more acidic isoperoxidases.     

Early anther ablation triggers parthenocarpic fruit development in tomato

Fruit set and fruit development in tomato is largely affected by changes in environmental conditions, therefore autonomous fruit set independent of fertilization is a highly desirable trait in tomato. Here, we report the production and characterization of male‐sterile transgenic plants that produce parthenocarpic fruits in two tomato cultivars (Micro‐Tom and Moneymaker). We generated male‐sterility using the cytotoxic gene barnase targeted to the anthers with the PsEND1 anther‐specific promoter. The ovaries of these plants grew in the absence of fertilization producing seedless, parthenocarpic fruits. Early anther ablation is essential to trigger the developing of the transgenic ovaries into fruits, in the absence of the signals usually generated during pollination and fertilization. Ovaries are fully functional and can be manually pollinated to obtain seeds. The transgenic plants obtained in the commercial cultivar Moneymaker show that the parthenocarpic development of the fruit does not have negative consequences in fruit quality. Throughout metabolomic analyses of the tomato fruits, we have identified two elite lines which showed increased levels of several health promoting metabolites and volatile compounds. Thus, early anther ablation can be considered a useful tool to promote fruit set and to obtain seedless and good quality fruits in tomato plants. These plants are also useful parental lines to be used in hybrid breeding approaches.     

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.     

The Solanum lycopersicum AUXIN RESPONSE FACTOR 7 (SlARF7) mediates cross-talk between auxin and gibberellin signalling during tomato fruit set and development

Transgenic tomato plants (Solanum lycopersicum L.) with reduced mRNA levels of AUXIN RESPONSE FACTOR 7 (SlARF7) form parthenocarpic fruits with morphological characteristics that seem to be the result of both increased auxin and gibberellin (GA) responses during fruit growth. This paper presents a more detailed analysis of these transgenic lines. Gene expression analysis of auxin-responsive genes show that SlARF7 may regulate only part of the auxin signalling pathway involved in tomato fruit set and development. Also, part of the GA signalling pathway was affected by the reduced levels of SlARF7 mRNA, as morphological and molecular analyses display similarities between GA-induced fruits and fruits formed by the RNAi SlARF7 lines. Nevertheless, the levels of GAs were strongly reduced compared with that in seeded fruits. These findings indicate that SlARF7 acts as a modifier of both auxin and gibberellin responses during tomato fruit set and development.     

The Effects of Salinity upon Galactosyl-Glycerol Content and Concentration of the Marine Red Alga Porphyra purpurea (Roth) C.Ag.

Changes in the major alcohol-soluble, low molecular weight carbohydratesof P. purpurea, O--D-galactopyranosyl-(1-2)-glycerol (‘floridoside’)and O--D-galactopyranosyl-(1-1)-glycerol (‘isofloridoside’),have been examined in response to salinity variation. ‘Floridoside’is shown to vary in absolute amount, increasing in hypersalineand decreasing in hyposaline media. ‘Isofloridoside’content per cell does not change in a similar manner. Responsesare almost identical under light or dark conditions, ‘floridoside’changes being complete within 24 h. Decreasing the externalwater potential using ionic and non-ionic solutes has the sameeffect upon galactosyl-glycerol content. The amount of ‘floridoside’synthesized, and degraded under hypersaline and hyposaline conditionsrespectively is shown to be insufficient to restore cell volumeto its original value. It is therefore suggested that the primaryfunction of ‘floridoside’ increases in concentratedsea-waters is that of a compatible solute, serving to protectthe cell during periods when the external salt content is increaseddramatically.