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Accessory fruit
Accessory fruit
Accessory fruit
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A selection of accessory fruits (from left to right: pear, fig, and strawberry)

An accessory fruit is a fruit that contains tissue derived from plant parts other than the ovary. In other words, the flesh of the fruit develops not from the floral ovary, but from some adjacent tissue exterior to the carpel (for example, from receptacles or sepal). As a general rule, the accessory fruit is a combination of several floral organs, including the ovary. In contrast, true fruit forms exclusively from the ovary of the flower.[1]

Accessory fruits are usually indehiscent, meaning that they do not split open to release seeds when they have reached maturity.[2]

Incorporated organs

[edit]
See also: Glossary of botanical terms

The following are examples of accessory fruits listed by the plant organ from which the accessory tissue is derived:[3]

Accessory Fruit Organs
Organ Fruit
Hypanthium apple, pear, rose hip
Involucre pineapple
Peduncle cashew apple
Perianth anthocarps of the Nyctaginaceae
Receptacle fig, mulberry, pineapple, strawberry
Calyx eastern teaberry, rose apple

Fruit with fleshy seeds, such as pomegranate or mamoncillo, are not considered to be accessory fruits.[4]

Examples

[edit]

Apples and pears

[edit]
Apple section, showing seeds plus papery expression of the ovary, surrounded by tissue formed from ripening of the hypanthium.

The part of apples and pears that is consumed is, in fact, the hypanthium. The ovary is the papery core that surrounds the apple seeds. As the hypanthium ripens it forms the edible tissues.[5]

Roses

[edit]

For roses, the hypanthium is the tissue that composes the edible part of rosehips. Roses and apples are both members of the Rosaceae family; the fact that they have similar fruit morphology is a major consideration in placing them in the same taxonomic family.[5]

Strawberries

[edit]
On this strawberry, the many pips located on the surface have germinated in a phenomenon known as vivipary. The pips of the strawberry are its true fruit.

The edible part of the strawberry is formed, as part of the ripening process, from the receptacle of the strawberry flower. The true fruits (hence, containing the seeds) are the roughly 200 pips (which are, technically, achenes, a true fruit that contains a single seed from a single ovary). These pips dot the exterior of the strawberry.[6]

Cashew apple

[edit]
The cashew 'apple' and its attached drupe, which contains the edible seed.

The cashew apple is an oval- or pear-shaped structure that develops from the pedicel and the receptacle of the cashew flower and is technically called a hypocarpium.[7][8][9] It ripens into a yellow or red structure about 5–11 cm (2–4+1⁄4 in) long.[10][8] The true fruit of the cashew tree is a kidney–shaped drupe that grows at the end of the cashew apple; the seed inside this drupe is the cashew nut of commerce.[10]

Pineapple

[edit]
Pineapple in flower

The pineapple is formed when 50 to 200 unpollinated flowers coalesce in a spiral arrangement— the flowers form individually and then fuse as a single 'multiple fruit'. The ovaries develop into berries and the fruit forms around an intercalary spike. The intercalary inflorescence (cluster of flowers) results when the terminal cluster of flowers are left behind by the growth of the main axis of the plant. Each polygonal area on the pineapple's surface is an individual flower.[11]

Research

[edit]

Current research has proposed that a single class of genes may be responsible for regulating accessory fruit formation and ripening.[12] A study using strawberries concluded that hormone signaling pathways involving gibberellic acid and auxin affect gene expression, and contribute to the initiation of accessory fruit development.[6] Metabolic modifications in different developing accessory fruit tissues are due to the varied distributions of compounds such as triterpenoids and steroids.[13]

See also

[edit]

References

[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Accessory fruit

View on Grokipedia
from Grokipedia
An accessory fruit, also known as a false fruit or pseudocarp, is a in which some or all of the fleshy portion develops from tissues outside the , such as the receptacle or , rather than solely from the pericarp ( wall). Unlike true fruits, which originate exclusively from the ripened following fertilization, accessory fruits incorporate additional floral structures that swell and mature alongside the -derived seeds. Accessory fruits play a crucial role in angiosperm by aiding through their attractive, often colorful and nutritious exteriors, which entice animals to consume and subsequently excrete the seeds. Common examples include the type, such as apples (Malus domestica) and pears (), where the fleshy forms the bulk of the surrounding a core of true fruit tissue; the strawberry ( × ananassa), an aggregate accessory fruit in which the enlarged receptacle bears numerous small achenes (true fruits) on its surface; and figs ( carica), where the fleshy receptacle encloses the structure. These fruits typically develop from perigynous or epigynous flowers, where the is positioned below or surrounded by other floral parts that contribute to the mature structure. In botanical classification, accessory fruits are distinguished from simple fruits (from a single ), aggregate fruits (from multiple ovaries in one flower), and multiple fruits (from fused ovaries of multiple flowers), though some, like strawberries, combine categories as aggregate accessory forms. They are generally indehiscent, meaning they do not split open at maturity to release seeds, relying instead on external agents for dispersal. This developmental strategy enhances the evolutionary success of many economically important crops, particularly in the family, by providing protective and appetizing packaging for the seeds.

Definition and Characteristics

Core Definition

An accessory fruit is a fruit in which some or all of the edible fleshy portion is derived from non-ovarian tissues, such as the receptacle, , or floral tube, rather than solely from the wall (pericarp). This distinguishes it from true fruits, which develop exclusively from the ripened and its associated structures. For instance, while a represents a true fruit with its fleshy portion originating from the ovarian pericarp, accessory fruits incorporate additional floral elements to form their edible parts. Key characteristics of accessory fruits include their typically fleshy texture and indehiscent nature, where the fruit does not split open at maturity to disperse , often resulting in a multi-layered composition from fused floral tissues. These fruits are adapted for animal dispersal through their appealing, non-ovarian , which protects and aids in the distribution of embedded . The term "pseudocarp" serves as a , emphasizing the inclusion of extraneous tissues beyond the true body. The concept of accessory fruits emerged in botanical classification during the late 19th century, with the term "accessory fruit" first recorded in 1858 to account for developmental contributions from non-ovarian parts in fruit morphology. This terminology arose as botanists refined distinctions between ovarian and extracarpellary tissues in angiosperm reproduction, building on earlier observations of fruit diversity in systematic studies.

Distinction from True Fruits

True fruits develop exclusively from the fertilized ovary(ies) of a flower, consisting solely of the pericarp (the ripened ovary wall) and any enclosed seeds. Simple fruits are true fruits that develop from a single pistil. In contrast, accessory fruits incorporate additional floral parts beyond the ovary, such as the receptacle or hypanthium, resulting in a composite structure where the edible portion is not derived primarily from the pericarp. For example, the tomato is a true fruit because it forms entirely from the ovary, while the apple is an accessory fruit, with its fleshy hypanthium surrounding the core derived from the ovary. Accessory fruits differ from other fruit categories like aggregate and multiple fruits in their developmental origins and structural dominance. Aggregate fruits arise from multiple ovaries (carpels) within a single flower that mature and fuse together, with the ovary tissues forming the primary structure, as seen in raspberries where individual drupelets cluster around a central receptacle. Accessory fruits, however, feature non-ovary tissues as the dominant component, distinguishing them from aggregates where ovaries predominate. Multiple fruits, formed from the fused ovaries of an entire (multiple flowers), often qualify as accessory when they include significant extrafloral tissues like the peduncle, such as in the . This highlights how accessory fruits emphasize accessory structures over pure ovarian development, unlike the ovary-centric true and aggregate forms.

Anatomical Components

Incorporated Plant Tissues

Accessory fruits incorporate various non-ovarian plant tissues that contribute to their structure and form the bulk of the mature fruit, distinguishing them from true fruits, which develop exclusively from the ovary wall. The primary tissues involved are the receptacle and hypanthium. The receptacle is the enlarged tip of the flower-bearing stem that supports the floral organs and expands to form fleshy or supportive portions of the fruit. In contrast, the hypanthium arises from the fusion of the basal portions of the sepals, petals, and stamens, creating a cup-shaped or tubular structure that surrounds the ovary and develops into the edible or protective layers of the fruit. Additional components may include the floral tube, an elongated extension of the receptacle or that forms a cylindrical structure contributing to the fruit's outer layers in certain cases. In aggregate or multiple accessory fruits, the pedicel (the stalk attaching the flower to the main axis) or rachis (the elongated axis of the ) can incorporate into the fruit body, providing a fleshy or structural base for clustered fruitlets. Rarely, bracts (modified leaves subtending the flower) or involucres (a whorl of fused bracts) integrate as protective or fleshy elements enclosing the true fruit structures. These incorporated tissues often form the flesh that encloses the true fruitlets, such as achenes or drupelets derived from the , thereby enhancing the 's size, texture, and dispersal capabilities. The layering typically positions the non-ovarian tissues externally, with the true fruit components embedded within, creating a composite unique to accessory fruits.

Structural Variations

Accessory fruits display a range of structural variations in the incorporation of non-ovarian tissues, such as the and receptacle, which contribute to their diverse morphologies. fruits develop from a swollen, cup-like structure formed by the fused bases of sepals, petals, and stamens, which envelops the and becomes the primary edible portion; for example, in pomes like apples, the fleshy surrounds the cartilaginous core containing the true fruitlets. Receptacular fruits, in contrast, arise from the enlargement of the flower's receptacle, the swollen basal portion that supports the floral organs, as seen in strawberries where this tissue expands into a juicy platform bearing embedded achenes (the true fruits). Composite types, such as the in figs, involve an inverted receptacle that forms a hollow, fleshy housing numerous tiny flowers on its inner surface, resulting in a unique internalized structure where the receptacle turns inside out during development. These variations contribute to morphological diversity in accessory fruits, particularly in the texture of the incorporated tissues and the number of contributing floral units. Accessory parts can be fleshy, providing succulence and moisture as in the of pears or the receptacle of mulberries, or dry and fibrous, such as the leathery outer layer in some hips where the accessory tissue aids in structural integrity rather than edibility. Simple accessory fruits originate from a single flower, integrating tissues from one set of floral organs around the , exemplified by the of a single apple blossom. In contrast, multiple accessory fruits form from fused structures of several flowers, as in the where multiple florets develop within the shared receptacle, or in pineapples where berries from adjacent flowers coalesce on an enlarged axis. Such structural adaptations serve key ecological functions, including enhanced protection of seeds and developing tissues, nutrient storage, and mechanisms. The enclosed , for instance, shields internal drupelets from environmental stressors and herbivores until maturity, while fleshy accessory tissues like the enlarged store carbohydrates and water to support establishment post-dispersal. These features also facilitate animal-mediated dispersal, with colorful, fleshy parts such as the red receptacular tissue in strawberries attracting birds that consume the and excrete viable seeds.

Developmental Biology

Formation Mechanisms

The formation of accessory fruits begins post-pollination, when non-ovarian tissues, such as the in fruits like apples, enlarge through successive phases of and cell expansion, while the fertilized develops into the true core housing . In this process, the accessory tissues contribute the bulk of the mature fruit's fleshy mass, distinguishing accessory fruits from those derived solely from the . For instance, in apples, initial growth after bloom relies exclusively on for approximately one week, doubling cell numbers exponentially, followed by a period of combined division and expansion for 3-4 additional weeks, after which expansion dominates to achieve final size. Similarly, in strawberries, the receptacle undergoes and expansion until about seven days after fall, transitioning thereafter to expansion alone. Developmental stages of accessory fruits encompass initiation, where flower clusters form in the prior ; floral development leading to bloom; and fruit set post-, marked by the resumption of in accessory tissues. During fruit set, typically cues the swelling of non-ovarian structures, though enables formation in certain cases, such as some varieties, without fertilization. In apples, fruit set involves followed by rapid tissue growth, with completing within 3-4 weeks post- to establish the fruit's cell count and potential size. These stages ensure the accessory tissues integrate with the ovarian core, forming a cohesive structure adapted for . Environmental factors play a key role in triggering and modulating tissue swelling during accessory fruit formation, with light, temperature, and cues acting as primary influencers. Adequate light exposure, particularly through open canopies during early post-bloom weeks, supports in apple by enhancing photosynthetic resources for growth. Temperature extremes can disrupt fruit set; for example, high temperatures during bloom impair success and subsequent expansion in strawberries, while optimal ranges (around 15-25°C) promote steady development. cues, including deposition and compatibility, are essential initiators, as incomplete leads to reduced tissue enlargement and uneven fruit formation across species like apples and strawberries.

Hormonal and Genetic Influences

The development of accessory fruits is profoundly influenced by plant hormones, particularly (GA), , and , which orchestrate , expansion, and maturation in non-ovarian tissues such as the receptacle or . promotes cell elongation and division in the receptacle, facilitating the enlargement of accessory structures during early fruit growth; for instance, in (an accessory fruit), the DELLA protein FveRGA1 represses and expansion, but post-fertilization GA signaling leads to its degradation, thereby activating growth in the receptacle. , often in synergy with GA, drives growth by regulating and differentiation in these tissues; studies in show that homeostasis genes dynamically adjust to support receptacle expansion, with levels peaking to stimulate non-ovarian tissue proliferation after . These hormones interact during , where modulates biosynthesis to coordinate softening and flavor development in the accessory portions, preventing premature while enabling climacteric-like responses in such as apple. At the genetic level, transcription factors play a central in specifying identity and regulating the incorporation of non-ovarian tissues into accessory fruits, particularly in the family. These genes, including SEPALLATA (SEP) homologs, coordinate the transition from floral organs to structures by activating downstream targets that promote or receptacle differentiation; in , for example, MIKC-type genes like PpMADS are differentially expressed during development to control tissue fusion and expansion. Research using model plants such as has elucidated mechanisms for non-ovarian tissue regulation, revealing how factors like SHATTERPROOF (SHP) and FRUITFULL (FUL) establish boundaries and promote valve-like growth in , providing insights applicable to accessory fruit patterning despite Arabidopsis's dry silique morphology. In accessory fruits, these genes ensure proper identity of receptacle-derived tissues, with altering shape and composition. Genetic variations, often arising from mutations in hormone-related or developmental genes, lead to abnormal accessory structures, including seedless varieties that rely on parthenocarpic growth of non-ovarian tissues. Mutations disrupting carpel or seed formation genes, such as those enhancing signaling, enable receptacle or expansion without fertilization; in apple, suppression of genes like MdMADS15 generates coreless, seedless fruits by redirecting growth to the hypanthium while maintaining overall accessory morphology. Similarly, parthenocarpic strawberry varieties, including some octoploid cultivars, develop enlarged receptacles without viable seeds through GA- and -induced growth, often propagated vegetatively for commercial production. These alterations highlight the plasticity of genetic networks in accessory fruits, where disruptions in or hormone receptor genes can yield viable, albeit structurally modified, forms without compromising edibility.

Prominent Examples

Rosaceae Family Fruits

The family is notable for producing a variety of accessory fruits, particularly pomes and hips, due to the characteristic inferior position in many subfamilies, which results in the fusion of floral parts with the during development. This anatomical feature leads to the enlargement of the —a cup-like structure formed from the fused bases of the calyx, corolla, and stamens—surrounding the true fruit tissues. In the subfamily Maloideae, apples (Malus domestica) and pears () exemplify pomes, where the fleshy exterior develops primarily from the , while the central core consists of papery carpels enclosing the seeds. The 's endocarp is cartilaginous and lined with sclereids, specialized sclerenchyma cells that contribute to the gritty texture, especially prominent in pears. These sclereids form clusters with thickened, lignified walls, enhancing structural integrity but altering during consumption. Rose hips, produced by species in the subfamily Rosoideae such as , represent another accessory fruit type, featuring an enlarged that encloses numerous achenes—the true dry fruits containing the seeds. These hips are particularly rich in , with contents varying by species and environmental factors; for instance, hips can contain up to 600 mg/L of ascorbic acid, supporting their traditional medicinal applications for immune support and anti-inflammatory effects.

Non-Rosaceae Examples

Accessory fruits outside the Rosaceae family exhibit diverse structural adaptations, often incorporating tissues from the receptacle, peduncle, or axis to form the bulk of the edible portion, while the true fruits—derived solely from the —remain subordinate. These examples highlight evolutionary variations in fruit development among monocots and other dicot families, contrasting with the hypanthium-dominated pomes typical of . The pineapple (Ananas comosus), from the Bromeliaceae family, represents a multiple accessory fruit formed through the coalescence of numerous individual flowers on a spike inflorescence. Each flower develops a berry (the true fruit), but the juicy, fibrous flesh primarily arises from the enlarged central axis and bracts, which fuse to create a composite structure with embedded seeds. This accessory tissue constitutes the majority of the fruit's mass, providing protection and aiding dispersal in tropical environments. In the family, the cashew apple (Anacardium occidentale) forms from the swollen peduncle and receptacle, known as the hypocarpium, which enlarges into a juicy, pear-shaped structure after fertilization. This accessory portion, often yellow to red and rich in , hangs below the true —a hard-shelled containing the edible kernel (nut). The pseudocarp-like apple swells dramatically, up to 5–10 cm long, while the drupe remains small and pendant, illustrating a unique inversion where the accessory tissue precedes the true fruit anatomically. Figs (Ficus carica), in the family, develop as syconia, a specialized multiple accessory fruit originating from a single inverted . The hollow, flask-shaped receptacle encloses hundreds of tiny flowers on its inner wall; upon maturation, the receptacle wall thickens into the edible, fleshy exterior, while the true fruits—small drupelets—form internally around the seeds. by fig wasps is essential for seed production in most varieties, with the accessory receptacle providing the bulk of the soft, sweet pulp that attracts dispersers. This structure, up to 5 cm in diameter, exemplifies how non-ovarian tissues can enclose and integrate multiple true fruits into a unified whole.

Significance and Applications

Economic Importance

Accessory fruits represent a significant portion of global agricultural output, with major crops such as apples and strawberries driving substantial economic activity through cultivation and trade. Global apple production reached approximately 84 million metric tons in the 2023/2024 marketing year, primarily led by , which accounts for over half of the total, followed by major producers like , the , and . production, another key accessory fruit, totaled over 9.5 million metric tons in 2022, with again dominating at around 3.4 million tons, supported by the , , and . Breeding programs for these crops focus on enhancing desirable traits like fruit size, yield, and flavor to meet market demands and improve economic viability; for instance, ongoing strawberry breeding efforts emphasize flavor improvement to boost consumer appeal and sales, while apple breeding incorporates ancient varieties for better size and resistance. Nutritionally, accessory fruits contribute to and economic value through their rich profiles of vitamins and , derived largely from the accessory tissues such as the receptacle in strawberries and the in apples. Strawberries are particularly noted for their high content, providing about 88 mg per cup of whole berries, which supports immune function and protection, making them a staple in health-focused markets. Pineapples, another accessory fruit, offer , a proteolytic with properties found in the stem and fruit, alongside at around 79 mg per cup, enhancing their appeal in functional foods and supplements. The accessory tissues in these fruits also provide significant —such as 4.5 grams per medium apple with skin and 3 grams per cup of strawberries—promoting digestive and adding to their commercial worth in wellness products. Commercially, accessory fruits underpin a diverse array of products that generate revenue in , beverage, and industries. Apples and are processed into juices, jams, and fresh markets, contributing billions to global trade; for example, the U.S. industry alone had a farm-gate value of approximately $3.4 billion in 2023. hips, the accessory fruit of roses, are utilized in jams, teas, and especially due to their high and content, with rosehip seed oil widely incorporated in skincare formulations for anti-aging and hydration benefits. In tropical regions, apples serve as a of nut production and are fermented into beverages like wine or feni, or processed into juices, adding economic value through local and emerging international markets, as seen in initiatives by companies like exploring apple juice.

Research Developments

Recent genetic studies have leveraged / technology to enhance accessory fruit tissue in strawberries, a classic example of accessory fruits where the edible portion derives primarily from receptacle tissue. Editing the polygalacturonase FaPG1 has produced strawberries with significantly improved firmness and slower post-harvest softening, reducing fungal susceptibility and extending shelf life. Similarly, knockout mutations in the auxin response factor ARF8 have increased fruit width and height, promoting larger accessory tissues and higher yields. These advancements, building on foundational hormonal influences, demonstrate CRISPR's precision in targeting developmental for commercial trait improvement. Post-2020 research has advanced understanding of hormone signaling pathways critical to accessory fruit formation, particularly interactions. In strawberries, knockdown of FveARF2 via accelerates ripening by elevating sugar and levels in the receptacle tissue, highlighting 's role in modulating accessory development. Broader studies on -GA crosstalk reveal how these hormones drive growth and in fruits like apples and peaches, with implications for resilient accessory structures. Such findings underscore evolving models of hormonal networks, informed by multi-omics approaches. Metabolic investigations in the 2020s have explored variations in and within accessory fruits, linking these to health benefits and consumer safety. Strawberry consumption enhances serum profiles and total capacity in adults with , attributed to high levels of polyphenols and in the accessory tissue. In apples, content varies by ; for example, 'Koksa Pomarańczowa' exhibits low Bet v 1 (4.24 µg/g) and levels, making it suitable for sufferers. These studies emphasize genotype-specific metabolic diversity, aiding breeding for varieties. Climate change research from the highlights heat stress impacts on accessory fruit development, with elevated temperatures disrupting physiological processes. In raspberries, a 4°C increase during fruit set reduces firmness, soluble solids, and content while increasing berry weight inconsistently, compromising quality. Strawberries under high temperatures show impaired , hormone dysregulation, and elevated , leading to smaller fruits and lower yields. These findings, from controlled heating experiments, stress the need for heat-tolerant cultivars. Despite progress, significant research gaps persist in accessory fruit biology. Evolutionary studies on wild accessory fruits remain limited, with most efforts focused on cultivated , leaving uncertainties in how shaped receptacle-dominated structures in non-domesticated lineages. For non-model like , while reference genomes exist, genomic maps of regulatory elements governing accessory tissue development are incomplete, hindering comparative analyses. Addressing these gaps could illuminate adaptive and enable broader genetic improvements.

References

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