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Parthenocarpy

Parthenocarpy is the development of fruit without fertilization of the ovules by pollen, resulting in the formation of seedless fruits.^[1]^[2] The term originates from Greek words meaning "virgin fruit," reflecting the absence of seed production.^[3] Parthenocarpy can occur naturally or be induced artificially, and it is classified into two main types: vegetative and stimulative.^[4] Vegetative parthenocarpy develops without any pollination stimulus and is obligate in certain species, such as bananas (Musa spp.) and pineapples (Ananas comosus), where fruits are consistently seedless.^[3]^[2] In contrast, stimulative parthenocarpy requires pollination to initiate fruit growth but proceeds without fertilization, as observed in grapes (Vitis vinifera).^[4] Facultative forms allow seed development if pollination occurs, providing flexibility in fruit set.^[2] This phenomenon holds significant agricultural value, as it enables reliable fruit production under environmental stresses like low pollinator activity or adverse weather, reduces labor for manual pollination, and yields seedless fruits with enhanced consumer appeal, improved flavor, and longer shelf life.^[2]^[3] In crops such as cucumbers, squash (Cucurbita spp.), and melons (Cucumis melo), parthenocarpy boosts yield stability and economic efficiency, particularly in greenhouse cultivation.^[2]

Definition and Fundamentals

Definition

Parthenocarpy, derived from the Greek words parthenos (virgin) and karpos (fruit), refers to the process by which fruits develop without the fertilization of ovules. The term was first introduced by the German botanist Fritz Noll in 1902 to describe fruit formation in the absence of pollination or seed development.^[5]^[6] In typical angiosperm reproduction, fruit development follows double fertilization, a unique process where one sperm cell fuses with the egg to form the diploid zygote (embryo), and a second sperm cell fuses with the central cell's polar nuclei to form the triploid endosperm. This endosperm provides nutritional support and generates hormonal signals that trigger the ovary to expand into a mature fruit, consisting of the pericarp—the wall derived from the ovary tissue—and the enclosed seeds. In parthenocarpy, however, no such fertilization occurs; instead, the ovary develops into a fruit autonomously, resulting in seedless fruits or those with rudimentary, non-viable seeds.^[7]^[8] Parthenocarpic fruits exhibit distinct characteristics compared to seeded fruits: they may exhibit reduced size in cases lacking full post-fertilization signals, but in obligate vegetative parthenocarpy, they often develop to comparable or larger edible portions without seeds. The seeds, if present, remain underdeveloped or entirely absent. Despite this, the pericarp often develops normally, forming the edible portion of the fruit, as hormonal signals—such as auxins and gibberellins—mimic the post-fertilization cues that promote cell division and elongation in the ovary wall. This process ensures fruit maturation without the need for seed formation, highlighting parthenocarpy's role as a deviation from standard reproductive pathways in flowering plants.^[9]^[10]

Historical Development

Early observations of seedless fruits, such as bananas (Musa spp.), date back to ancient cultivation practices in Southeast Asia around 7,000 years ago, where parthenocarpic mutants were selected for their desirable traits, though scientific recognition emerged in the 18th and 19th centuries through botanical descriptions of naturally occurring seedless varieties in crops like figs and grapes.^[11] Systematic studies began in the early 20th century among plant breeders, who documented parthenocarpic fruit development in response to environmental factors or genetic variations without pollination.^[12] The term "parthenocarpy" was coined in 1902 by German botanist Fritz Noll, who described the phenomenon in cucumbers (Cucumis sativus) as fruit formation without prior fertilization, drawing an analogy to parthenogenesis in animals.^[13] In the 1930s, H.B. Tukey advanced understanding through experiments on fruit set in strawberries (Fragaria × ananassa), showing that applications of growth-promoting substances could stimulate parthenocarpic development, laying groundwork for hormonal interventions.^[14] Felix G. Gustafson further contributed in 1936 by demonstrating that extracts from maturing fruits, later identified as gibberellins, could induce parthenocarpy in various species, highlighting the role of plant growth regulators. Post-World War II research accelerated with practical applications of synthetic auxins and gibberellins for inducing seedless fruits in horticulture. In the 1950s and 1960s, studies shifted toward biochemical elucidation, identifying auxins and gibberellins as key hormones that bypass pollination signals to promote ovary growth and fruit enlargement.^[15] Subsequent decades saw advances in molecular biology, with genetic engineering approaches in the 1990s and 2000s targeting genes like those encoding auxin biosynthesis enzymes to induce parthenocarpy in crops such as tomatoes. As of 2025, recent research has identified specific loci, such as THERMOSENSITIVE PARTHENOCARPY 4 in tomatoes, revealing feedback loops involving gibberellins and transcription factors that regulate fruit set under stress conditions.^[16]^[17]

Types and Mechanisms

Classification of Types

Parthenocarpy is classified into primary types based on the involvement of pollination and fertilization processes, with vegetative parthenocarpy defined as the autonomous development of fruit from an unfertilized ovary without any pollination stimulus.^[4] In this type, the ovary progresses to fruit formation independently, relying on internal developmental cues rather than external triggers.^[4] Conversely, stimulative parthenocarpy occurs when pollination provides a stimulus for fruit initiation, but fertilization does not take place, often involving pollen tube growth that fails to result in sperm-egg fusion.^[4] The criteria for differentiating these types center on the role of pollination: vegetative parthenocarpy requires none, while stimulative parthenocarpy depends on it as a trigger without completing fertilization.^[4] These distinctions ensure that classifications reflect both the physiological processes and practical implications for fruit development reliability.^[4] Obligate parthenocarpy, a subtype of vegetative parthenocarpy, occurs invariably without pollination in certain species, resulting in consistently seedless fruits, while facultative parthenocarpy manifests conditionally, allowing seed development if pollination and fertilization occur under favorable conditions.^[2] Obligate forms represent a fixed trait, often genetically determined for perpetual seedlessness, while facultative variants allow for sexual reproduction under favorable conditions, highlighting the spectrum of parthenocarpic expression.^[2] Hormonal signals, such as auxins and gibberellins, may underpin these variants but are modulated differently across types.^[2]

Biological Mechanisms

Parthenocarpy is primarily driven by hormonal signals that mimic the fertilization-induced activation of ovary development, with auxins (indole-3-acetic acid, IAA) and gibberellins (GA) playing central roles in initiating cell division and expansion in the ovary walls. Auxins promote pericarp growth by activating downstream signaling pathways, such as the ARF7/IAA9 complex, where repression of IAA9 leads to enhanced auxin responsiveness and ovary enlargement independent of pollination. In tomato, for instance, higher endogenous IAA levels in parthenocarpic ovaries facilitate seedless fruit formation by upregulating genes involved in cell proliferation. Gibberellins, particularly GA3, similarly trigger fruit set by binding to the GID1 receptor, which promotes the ubiquitination and degradation of DELLA proteins—key repressors of GA signaling—thereby derepressing growth-related transcription factors and mimicking the post-fertilization hormonal surge.^[18] The interplay between auxins and gibberellins forms feedback loops that sustain parthenocarpic development; auxin application or increased biosynthesis upregulates GA biosynthetic genes like GA20ox1, amplifying GA levels and further promoting cell division without seed initiation, while GA does not directly feedback to auxin levels in early stages. This hormonal cross-talk ensures coordinated pericarp expansion, as seen in transcriptome analyses of parthenocarpic tomato fruits where auxin-induced GA signaling dominates early fruit set. Cytokinins can enhance this loop by boosting both IAA and GA3 accumulation, though their role is auxiliary to the primary auxin-GA axis.^[18] Genetic factors contribute to parthenocarpy by disrupting seed formation pathways while preserving pericarp growth signals. In tomato, mutations in the MADS-box transcription factor gene SlAGL6 (also known as Pat-k) result in facultative parthenocarpy, where loss-of-function alleles prevent normal ovule development and seed set but allow hormone-driven ovary wall expansion, leading to viable seedless fruits without altering overall fruit morphology. Similarly, in strawberry, mutations in the DELLA-encoding gene FveRGA1 derepress GA signaling in the receptacle, enabling parthenocarpic fruit initiation and growth in the absence of fertilization by overriding repression of auxin-responsive genes, thus promoting larger, seedless accessory fruits. These genetic alterations highlight how targeted disruptions in reproductive versus vegetative development pathways can uncouple fruit set from seed production. Recent studies as of 2025 have identified additional regulatory mechanisms, such as the TSP4 locus in tomato, comprising SlIAA9 and SlANT genes that form a positive feedback loop to control thermosensitive parthenocarpy under heat stress.^[19]^[20]^[16] Environmental triggers, such as low temperatures and specific photoperiods, induce stimulative parthenocarpy by modulating hormone balances, particularly through ethylene regulation. Low nightly temperatures elevate auxin content in ovaries, as observed in cucumber and zucchini, promoting unpollinated fruit set via enhanced cell division without requiring genetic changes. Short-day photoperiods similarly boost auxin activity, facilitating parthenocarpic development in cucurbits by altering light-sensitive hormone biosynthesis. Ethylene modulation acts as a key integrator, where its inhibition—often triggered by these environmental cues—prevents ovary senescence and allows GA and auxin dominance, thereby enabling fruit growth in stimulative types under suboptimal conditions.^[2] At the molecular level, parthenocarpy-specific genes like the pat (SlIAA9) in tomato influence auxin transport and signaling, where mutations cause constitutive auxin responses by repressing negative regulators, leading to parthenocarpic fruit via feedback amplification of auxin efflux carriers such as PIN proteins. This creates a self-reinforcing loop: increased auxin transport to the ovary upregulates local biosynthesis and signaling, sustaining pericarp development independent of fertilization signals, as evidenced in auxin transport inhibitor studies that induce seedless fruits through GA mediation.^[21]

Natural and Induced Examples

Naturally Occurring Cases

Parthenocarpy occurs naturally in various wild and cultivated plants, enabling fruit development without fertilization and often serving as an adaptation to environmental challenges such as limited pollinators. This phenomenon is documented in numerous angiosperm taxa, with higher prevalence in polyploid species (52.1% of parthenocarpic taxa are polyploid, compared to 34.5% in angiosperms overall), where it frequently links to sterility or self-incompatibility mechanisms that prevent successful seed formation.^[22] In natural settings, such fruits are typically viable for dispersal but rely on vegetative propagation for species continuation, as seen in isolated populations where pollination is unreliable.^[22] A prominent example is the banana (Musa spp.), particularly triploid varieties like the Cavendish, which exhibit natural vegetative parthenocarpy due to polyploidy-induced sterility, resulting in seedless fruits that enhance dispersal efficiency in tropical habitats.^[23] Similarly, in figs (Ficus carica), common varieties such as Brown Turkey display obligatory parthenocarpy, producing seedless syconia without pollination or fig wasps, an adaptation that bypasses self-incompatibility and supports reproduction in regions lacking specific pollinators.^[24] This trait allows the breba crop to form independently, contributing to the plant's evolutionary success in Mediterranean and temperate wild populations.^[24] Pineapple (Ananas comosus) represents an obligate parthenocarpic species, consistently developing seedless fruits in wild relatives, which facilitates vegetative spread via crowns or slips in isolated or bromeliad-dominated ecosystems.^[2] In the Cucurbitaceae family, facultative parthenocarpy appears in cucumber (Cucumis sativus) germplasm, where unpollinated ovaries can develop into seedless fruits under stress like low temperatures, linked to hormonal changes mimicking fertilization signals.^[2]^[22] Evolutionarily, natural parthenocarpy often integrates with apomictic reproduction, providing an advantage in seedless fruit production for dispersal in pollinator-scarce or isolated populations, as it deceives seed predators while promoting clonal propagation.^[22] This sporadic occurrence across angiosperm families underscores its role in enhancing reproductive assurance without genetic recombination, particularly in polyploid lineages facing self-incompatibility barriers.^[22]

Artificially Induced Cases

Artificially induced parthenocarpy involves human interventions to promote seedless fruit development without fertilization, primarily through hormonal applications, physical manipulations, and genetic modifications. Hormonal methods dominate agricultural practices, utilizing plant growth regulators to mimic pollination signals and stimulate ovary growth. Common regulators include gibberellic acid (GA3), applied at concentrations typically ranging from 100 to 200 ppm, and auxins such as 2,4-dichlorophenoxyacetic acid (2,4-D) or naphthaleneacetic acid (NAA) at 20 to 50 ppm, often sprayed on flowers or young fruits.^[25]^[26] These treatments are timed for pre-bloom or anthesis stages to maximize efficacy, as earlier or later applications may reduce fruit set or cause deformities.^[27] In seedless grape production (Vitis vinifera), GA3 sprays have been standard since the late 1950s, following initial tests in 1957 that demonstrated increased berry size and reduced seed formation when applied at bloom (40-80% cap-off) or berry set (5-10 mm diameter). Concentrations of 100 ppm pre- and post-bloom, combined with multiple applications, yield elongated clusters and larger, seedless berries, transforming varieties like Thompson Seedless into commercial staples by 1962.^[28]^[29] For tomatoes (Solanum lycopersicum), especially in greenhouse settings, GA3 or auxin mixtures (e.g., 30-40 ppm GA3) applied to unpollinated ovaries induce parthenocarpic fruit set, bypassing pollination challenges in controlled environments and producing uniform, seedless fruits.^[30]^[18] In cucumbers (Cucumis sativus), auxin or brassinosteroid analogs like 24-epibrassinolide (0.2 μM) achieve high parthenocarpic rates, often exceeding 80% fruit set without seeds, though variable environmental uptake can lead to uneven sizing or misshapen fruits.^[26]^[31] Physical techniques, such as emasculation to remove anthers before anthesis or pollination with gamma-irradiated (infertile) pollen, provide non-chemical induction by preventing fertilization while delivering stimulatory signals from pollen tubes. These methods are labor-intensive but effective in experimental settings for crops like tomatoes, yielding parthenocarpic fruits comparable to hormonal treatments without residue concerns.^[32]^[13] Modern genetic approaches offer stable induction without repeated applications. For instance, insertion of the DefH9-iaaM chimeric gene, which drives ovule-specific auxin biosynthesis from tryptophan, produces parthenocarpic eggplants (Solanum melongena) with enlarged, seedless fruits under diverse conditions, including suboptimal pollination environments; this transgene has enhanced winter yields by up to 33% in protected cultivation.^[33]^[34] Similar engineering in tomatoes and cucumbers confirms the iaaM gene's role in deregulating hormonal balance for reliable parthenocarpy.^[35] Despite successes, challenges like inconsistent hormone distribution and potential fruit abnormalities persist, necessitating optimized protocols for commercial scalability.^[25]

Significance and Applications

Ecological Role

Parthenocarpy enables plants to produce fruits without pollination and fertilization, providing a reproductive advantage in environments where pollinators are scarce or unreliable, such as isolated habitats or during periods of adverse weather that hinder insect activity.^[36] This mechanism ensures fruit development and potential seed dispersal even under low pollination success, promoting plant persistence in unstable or disturbed ecosystems.^[37] In natural settings, parthenocarpy contributes to biodiversity by enhancing the survival and spread of certain species, particularly weedy or invasive ones. For instance, in wild parsnip (Pastinaca sativa), parthenocarpic fruits serve as decoys that divert seed predators like flies away from seeded fruits, thereby protecting viable offspring and facilitating establishment in competitive environments.^[38] Similarly, in tropical dry forests, species such as Bursera morelensis produce parthenocarpic fruits that mimic seeded ones in appearance and abundance, attracting frugivorous birds for dispersal while reducing predation by granivores; experimental observations showed that trees with higher proportions of seedless fruits (up to 52%) experienced increased bird visitation and lower seed loss.^[39] This deceit strategy occurs in several plurispermic tropical species, where parthenocarpy supports fruit set under variable pollination conditions and aids in mimicking seed dispersal cues to maintain population dynamics.^[37] Parthenocarpy influences ecosystem interactions by decreasing plants' dependence on pollinators for fruit production, which may alleviate pressure on declining insect populations in pollinator-limited habitats.^[36] However, since parthenocarpic plants typically still produce flowers offering nectar and pollen, they can continue supporting pollinator foraging without fully eliminating mutualistic ties. In some perennials, the absence of seeds in parthenocarpic fruits limits contributions to the soil seed bank, potentially altering long-term community regeneration and favoring clonal propagation over sexual recruitment in dynamic landscapes. For example, naturally parthenocarpic bananas (Musa spp.) rely on vegetative spread, which sustains populations in fragmented tropical habitats but restricts seedling establishment.^[40] Evolutionarily, parthenocarpy offers reproductive assurance but involves trade-offs, such as reduced genetic diversity when paired with clonal propagation, as seen in polyploid parthenocarpic lineages where hybridization disrupts sexual reproduction and promotes uniformity.^[37] This can enhance short-term survival in harsh conditions but limits adaptability to changing environments compared to sexually reproducing counterparts.^[40]

Agricultural and Commercial Uses

Parthenocarpy has revolutionized crop production by enabling the development of seedless varieties that simplify processing and enhance market appeal in agriculture. In bananas and cucumbers, for instance, parthenocarpic cultivars ensure consistent fruit set without pollination, facilitating easier consumption and reducing waste.^[3] Similarly, in strawberries, genetic engineering to induce parthenocarpy has increased fruit size by 20% to over 100% in transgenic lines of Fragaria vesca, allowing reliable yields in controlled environments or short growing seasons where pollination is limited.^[41] These improvements stem from breeding strategies focused on facultative parthenocarpy, where fruit set occurs with or without pollination, as seen in hybrid selections for cucurbits like cucumbers.^[2] Commercial adoption began notably in the 1930s with the introduction of the parthenocarpic cucumber cultivar 'Geneva' in the United States, bred for consistent fruit set in greenhouses without manual pollination, thereby cutting labor costs.^[42] The global market for seedless fruits, including those produced via parthenocarpy, is projected to grow significantly, driven by consumer demand for convenient, ready-to-eat produce that minimizes seed-related processing and disposal.^[43] Parthenocarpic varieties also boost yields under suboptimal conditions; for example, in cucurbits, they achieve fruit set rates over 50% in selected germplasms, enhancing economic viability for growers facing pollinator shortages.^[2] Key benefits include heightened consumer preference for seedless fruits, which command premium prices due to their convenience, and reduced post-harvest waste from seed removal.^[44] However, challenges persist, such as lower plant vigor in some parthenocarpic lines, which may necessitate additional supports or hybrid vigor enhancements to maintain productivity.^[45] Ongoing research targets climate-resilient induction methods, including hormone applications and genetic tools like QTL mapping. Recent advances as of 2025 include the identification of the THERMOSENSITIVE PARTHENOCARPY 4 gene in tomatoes, which optimizes auxin signaling for parthenocarpy under heat stress, and PbCYP78A6 in pears for induced pseudo-embryo development, promoting stable yields for crops like tomatoes and cucurbits amid environmental pressures.^[2]^[16]^[46]

Distinctions from Similar Phenomena

Parthenocarpy differs fundamentally from parthenogenesis, as the former involves the development of fruit tissue in the absence of fertilization without any embryo formation, resulting in seedless or rudimentary seed structures, whereas parthenogenesis entails the asexual development of an embryo from an unfertilized egg cell, often leading to viable offspring. This distinction is evident in examples like seedless bananas, where parthenocarpy produces fruit without embryos, in contrast to parthenogenetic reproduction in aphids, which generates clonal embryos capable of further development.^[22] In comparison to apomixis, parthenocarpy focuses solely on fruit maturation without viable seeds, bypassing both fertilization and embryo development, while apomixis is an asexual seed-forming process that incorporates parthenogenesis to produce unreduced, clonal seeds that can germinate.^[22] For instance, apomictic species like Tripsacum dactyloides yield seeds genetically identical to the parent plant through mechanisms such as apospory or diplospory, whereas parthenocarpic fruits in tomatoes or grapes contain no functional seeds, emphasizing fruit tissue expansion over seed production.^[47] Parthenocarpy also contrasts with vivipary, where the latter involves premature seed germination within the fruit while still attached to the parent plant, often as an adaptation to harsh environments, leading to the direct emergence of plantlets; in parthenocarpy, however, fruits develop without fertilization and typically feature non-viable or absent seeds that do not germinate.^[48] Vivipary is observed in mangroves like Rhizophora species, where seedlings sprout viviparously for immediate establishment, unlike the inert, seedless fruits from parthenocarpic processes in crops such as cucumbers.^[49] A key differentiator of parthenocarpy is its restriction to angiosperms, where it pertains specifically to the ovary-derived fruit structures, whereas parthenogenesis and apomixis occur across diverse taxa including animals and non-angiosperm plants, and vivipary manifests in both seed-producing and non-seed plants as a germination strategy.^[22]

Common Misconceptions

One prevalent misconception is that all seedless fruits result from genetic modification, when in fact most arise from natural or traditionally bred parthenocarpic processes predating modern biotechnology.^[3] For instance, cultivated bananas (Musa spp.) exhibit natural parthenocarpy due to their sterility, producing seedless fruits without genetic engineering, a trait selected by humans over millennia from wild ancestors.^[23] Genetically modified seedless varieties remain rare in commercial production, with parthenocarpy achieved primarily through selective breeding or hormonal induction.^[50] Another common error is the belief that parthenocarpy eliminates any need for pollination, whereas certain types, known as stimulative parthenocarpy, require a pollination stimulus to trigger fruit development, though fertilization does not occur.^[4] In such cases, like certain grape varieties, pollen deposition initiates hormonal signals for fruit set without seed formation, distinguishing it from vegetative parthenocarpy, which proceeds autonomously without external cues.^[51] This nuance highlights that parthenocarpy does not universally bypass pollinators but can enhance fruit production under suboptimal pollination conditions. It is also mistakenly assumed that parthenocarpic fruits are inherently inferior in quality to their seeded counterparts, yet studies show they often exhibit desirable traits such as higher soluble solids content, leading to sweeter flavors, though they may be smaller or less firm and sometimes require pollination for optimal yield. For example, in tomatoes, parthenocarpic fruits display elevated sugar levels compared to seeded ones, improving palatability while reducing seed-related waste.^[52] These qualities make them valuable in agriculture, countering notions of overall inferiority. Such misconceptions often stem from media sensationalism surrounding genetically modified crops, which amplifies fears of "unnatural" interventions, and confusion with hybrid sterility, where seedlessness arises from chromosomal incompatibilities rather than targeted parthenocarpy.^[53] This hype overlooks the long history of natural seedless varieties, fostering undue skepticism toward parthenocarpic produce.^[3]

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Publication : USDA ARS
Jul 1, 2007 · The lesser vigor (in many cases) of parthenocarpic clones suggests that this germplasm will be most useful, initially, in improving fruit ...
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[PDF] Improvement of apomictic seed and parthenocarpic fruit ...
Apomixis and parthenocarpy are broadly botanically defined as embryo and seed, and fruit development respectively, in the absence of fertilization.
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METHYLTRANSFERASE1 and Ripening Modulate Vivipary during ...
Vivipary, wherein seeds germinate prior to dispersal while still associated with the maternal plant, is an adaptation to extreme environments. It is normally ...
[47]
No, there's nothing wrong with seedless fruits | New Scientist
Sep 8, 2021 · Seedless fruits are the result of a biological process called parthenocarpy – the development of a fruit without prior fertilisation. While ...<|separator|>
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Parthenocarpic fruit quality and production under pollinator ...
Mar 15, 2024 · Non-pollinated parthenocarpic fruits took longer to develop and tended to be smaller, sweeter, and less firm than open-pollinated fruits.
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[PDF] Effects of Parthenocarpy on Fruit Quality in Tomato - ASHS Journals
Parthenocarpic fruit were found to have higher percent soluble solids than low-seeded fruit, which, in turn, had higher values than seeded fruit. In general,.
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Misinformation in the media: global coverage of GMOs 2019-2021
Likewise, there is a clearly stated consensus among major national and international scientific bodies that food derived from GM crops is as safe as any other.