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Multiple fruit

A multiple fruit, also known as a collective or infructescence fruit, is a composite fruit in botany that forms when the ovaries of multiple flowers within an inflorescence merge together during maturation, creating a single fruit structure that incorporates the ripened ovaries, receptacles, and sometimes other floral parts from each flower.^[1] This fusion results in a fruit composed of numerous individual fruitlets, each derived from a separate flower, enclosing seeds that aid in plant reproduction and dispersal.^[2] Unlike simple fruits, which develop from a single ovary (such as a peach), or aggregate fruits, which arise from multiple ovaries within one flower (such as a raspberry), multiple fruits originate from an entire cluster of flowers packed on a common axis.^[3] Prominent examples include the pineapple (Ananas comosus), where the edible fruit develops from the fused ovaries of 100 or more flowers in a spike inflorescence, producing its characteristic segmented, juicy exterior; the mulberry (Morus spp.), resembling a blackberry but formed from drupelets of separate flowers; and the jackfruit (Artocarpus heterophyllus), the largest tree-borne fruit, aggregating numerous florets into a massive, fibrous mass.^[4]^[2] These fruits are typically fleshy and often brightly colored or sweetly scented to attract animal dispersers, contributing to their ecological roles and economic value in tropical and subtropical agriculture.^[5] The pineapple, for instance, is a major global crop, while mulberry trees provide leaves for silkworm cultivation and jackfruit provides versatile nutrition in Southeast Asia.^[6]

Definition and Characteristics

Definition

A multiple fruit, also known as a composite fruit or collective fruit, is a type of fruiting body that develops from the fused ovaries of multiple flowers arranged in an inflorescence, resulting in a single cohesive structure often referred to as an infructescence.^[1]^[7]^[8] This botanical category is distinguished from other fruit types by its origin from the collective maturation of several distinct flowers rather than from a single flower's ovary or multiple carpels within one flower.^[9]^[10] The concept of multiple fruits emerged in 19th-century botanical systematics, with early definitions appearing in English literature through John Lindley's 1832 Introduction to Botany and related classifications by Alphonse de Candolle, who contributed to clarifying fruit terminology amid confusions with aggregate forms.^[11]

Key Characteristics

Multiple fruits are distinguished by their composite structure, arising from the ovaries of multiple flowers within an inflorescence that fuse together to form a single fruiting body. This fusion involves the maturation and coalescence of numerous individual ovaries, creating a unified mass where the boundaries between original floral units may become indistinct.^[12]^[13] A hallmark of multiple fruits is the frequent incorporation of persistent floral parts, such as sepals, bracts, or receptacles, which integrate into the fruit wall and contribute to its overall architecture. These elements often provide structural support or additional tissue layers, enhancing the fruit's cohesion. Externally, multiple fruits typically exhibit irregular shapes due to the variable arrangement and development of the constituent flowers, along with embedded seed-like structures derived from the individual ovaries, which may appear as distinct units within the composite. Texture variations range from fleshy, often juicy forms to drier types, depending on the degree of tissue elaboration in the fused components.^[14]^[12] The pericarp in multiple fruits is uniquely composed from fused tissues of multiple origins, including the walls of numerous ovaries as well as accessory floral structures like perianth or rachis. This results in a heterogeneous outer layer that lacks the uniform derivation seen in simple fruits, with the exocarp, mesocarp, and endocarp elements blending across the diverse sources to form a protective and sometimes edible envelope.^[13]^[14]

Formation and Development

Developmental Process

The developmental process of multiple fruits begins with the initiation of an inflorescence, a cluster of closely spaced flowers on a single axis, where pollination occurs across multiple florets. Successful pollination leads to fertilization of the ovules within each flower's ovary, triggering the enlargement of these ovaries as they begin to develop into individual fruitlets. This post-fertilization phase is stimulated by hormones produced by the developing seeds, which diffuse into the ovary walls and promote initial growth through cell division.^[15]^[16] As development progresses, the fusion process integrates these individual fruitlets into a cohesive structure. The enlarging ovaries, along with surrounding tissues such as the receptacles, peduncles, and other floral parts like bracts and perianths, merge through continued cell expansion and tissue intergrowth. This coalescence occurs as the fruitlets ripen, resulting in a single, unified fruit body that incorporates the collective ovaries from the entire inflorescence. The fused structure is a key characteristic, distinguishing multiple fruits from other types.^[5]^[12] The timeline of multiple fruit development encompasses several sequential stages: flowering and pollination, followed by ovary maturation involving rapid cell division and expansion, and culminating in ripening. During ovary maturation, auxins play a central role in regulating cell expansion, while gibberellins contribute to overall growth coordination across the inflorescence. Ripening marks the final phase, where hormonal shifts, including increased ethylene, induce changes in texture, color, and flavor to facilitate seed dispersal, completing the transformation into a mature multiple fruit.^[17]^[16]

Anatomical Features

Multiple fruits display a distinctive composite anatomy arising from the coalescence of ovaries, receptacles, and associated floral tissues from numerous flowers in an inflorescence. The exocarp, or outer skin, forms from the fused epidermal and subepidermal layers of the individual fruitlets, creating a unified protective covering that varies from tough and waxy in species like the pineapple (Ananas comosus) to thin and leathery in the fig (Ficus carica).^[18]^[19]^[20] This layer often includes cuticularized cells to minimize water loss and deter pathogens. The mesocarp, the fleshy middle region, develops primarily from the expanded receptacles and fused mesocarp tissues of the original flowers, resulting in a parenchymatous matrix rich in storage cells that imparts succulence and nutritional value. In the pineapple, this layer consists of juicy, fibrous parenchyma interspersed with raphides-containing idioblasts, while in the fig syconium, it forms the whitish, edible interior rind derived from receptacle tissue.^[21]^[19]^[20] The endocarp, surrounding the seeds, originates from the innermost walls of the individual ovaries and often remains distinct for each seed cavity, providing localized enclosure. In multiple fruits, these endocarps can be thin and membranous or hardened with stone-like cells, as seen in the individual drupelets of the fig, where each contains a single seed encased in a lignified endocarp. Sclerenchyma tissues, including fibers and sclereids, are prevalent in the endocarp and intervening septa, enhancing mechanical strength and seed protection; for instance, sclerenchymatous fibers reinforce the core of the pineapple, while sclereids occur in the fig's drupelet walls.^[21]^[20]^[19] A hallmark of multiple fruit anatomy is the presence of numerous locules or seed cavities, each corresponding to the ovary of an original flower, separated by thin parenchymatous or fibrous partitions. These locules house one or more seeds per cavity, with the overall arrangement reflecting the inflorescence pattern—hundreds in the pineapple's spirally arranged fruitlets and similarly numerous in the fig's enclosed syconium. Vascular tissue integration is intricate, as the phloem and xylem bundles from multiple flowers anastomose with those of the central axis, forming a reticulate network that ensures efficient nutrient and water distribution across the fused structure; in the pineapple, these bundles are prominently arranged in the fibrous core, while in the fig, they ramify through the receptacle wall to supply the embedded drupelets.^[19]^[20]^[21] These features emerge through the developmental fusion of floral parts, yielding a structurally unified yet internally diverse organ.^[18]

Types and Examples

Common Examples

One prominent example of a multiple fruit is the pineapple (Ananas comosus), a syncarp formed by the fusion of approximately 100 to 200 individual flowers into a single composite structure.^[22] Each hexagonal "eye" on the pineapple's surface corresponds to an individual berry derived from one of these flowers.^[23] Native to the Paraná–Paraguay River basin in South America, the pineapple is widely cultivated in tropical regions for its sweet, juicy, edible fruit, which is consumed fresh, juiced, or processed.^[24]^[25] Another well-known multiple fruit is the fig (Ficus carica), classified botanically as a syconium that develops from a single inverted inflorescence enclosing hundreds of tiny flowers.^[26] Inside this fleshy, hollow receptacle, the flowers mature into numerous small drupelets, which collectively form the edible portion of the fruit.^[27] Originating from the Mediterranean region to Central Asia, figs are extensively cultivated in warm climates worldwide for their sweet, nutritious fruits, often eaten fresh, dried, or in preserves.^[28]^[29] The mulberry (Morus spp.), such as the white mulberry (M. alba) native to East Asia and the red mulberry (M. rubra) native to eastern North America, exemplifies a sorosis formed from a catkin-like inflorescence where multiple flowers develop into a cohesive, fleshy cluster.^[30]^[31]^[32] This aggregate-like structure consists of numerous tiny drupelets that create a juicy, elongated fruit resembling a elongated blackberry in appearance.^[33] Mulberries are edible and valued for their sweet-tart flavor, with species cultivated globally for fresh consumption, silkworm feed, and in jams or wines.^[34]

Variations and Types

Multiple fruits exhibit structural variations primarily based on the type of inflorescence from which they develop and the nature of the fused floral parts. One major subtype is the sorosis, formed from a spike, spadix, or catkin inflorescence where the fleshy axis, bracts, and perianths consolidate with the ovaries to create a unified structure.^[35] Examples include the mulberry (Morus spp.), which develops from a catkin-like inflorescence, resulting in a collective drupe-like fruit.^[36] Another subtype is the syconium, derived from a hypanthodium inflorescence in which numerous tiny flowers are enclosed within a fleshy, hollow receptacle that inverts during development.^[37] The common fig (Ficus carica) exemplifies this, with its syconium appearing as a single fruit but comprising hundreds of florets.^[38] The coenocarpium represents a specialized subtype similar to the sorosis, formed by the consolidation of ovaries and other floral parts from multiple flowers on an elongated or fleshy axis.^[36] Jackfruit (Artocarpus heterophyllus) illustrates this variation, where the massive fruit arises from a syncarpous inflorescence of numerous female flowers, forming a syncarp that can weigh up to 55 kg.^[39] In contrast to the more familiar pineapple, these subtypes highlight morphological diversity in how receptacles and ovaries integrate.^[40] Multiple fruits also vary between fleshy and dry forms, influencing their ecological roles and edibility. Fleshy multiple fruits, such as those in the sorosis and syconium subtypes, typically feature juicy pericarps that aid animal dispersal, as seen in figs and mulberries.^[37] Dry multiple fruits, however, develop hardened or fibrous tissues with minimal succulence, often serving wind or mechanical dispersal. The Osage orange (Maclura pomifera), a multiple fruit composed of fused druplets on a central axis, exemplifies this dry form; its bumpy, yellow-green syncarp structure, up to 15 cm in diameter, contains sticky latex but lacks palatable flesh.^[41] Similarly, breadfruit (Artocarpus altilis), while fleshy when immature, matures into a starchy, less juicy multiple fruit (syncarp) that can be considered in its dry-utilization phase for food processing, contrasting with the perpetually succulent jackfruit.^[42] Less common examples underscore further diversity, particularly in tropical species. Banyan figs (Ficus benghalensis), like other Ficus species, produce syconia but on a grander scale due to the tree's expansive growth; these multiple fruits, pollinated by specific wasps, vary in size and ostiole structure to accommodate hemi-epiphytic habits.^[38] Tropical multiple fruits occasionally exhibit aggregate-like appearances, such as in certain Moraceae, where fused inflorescences create irregular, bulbous forms adapted to diverse habitats.^[36] These variations reflect adaptations in inflorescence architecture and tissue fusion across angiosperm lineages.^[43]

Comparison to Other Fruits

With Simple Fruits

Simple fruits develop from the ovary of a single flower, which may consist of one carpel or multiple fused carpels, resulting in a structure containing seeds from a single reproductive unit.^[44] For instance, the apple is a pome, a type of simple fleshy fruit where the edible portion derives primarily from the receptacle and floral tube surrounding the ovary, while the pea pod represents a simple dry fruit known as a legume, formed from a single carpel that splits along two seams to release seeds.^[45] In contrast, multiple fruits arise from the fused ovaries of an entire inflorescence, incorporating contributions from numerous flowers.^[46] Key differences between simple and multiple fruits lie in their origins, structural composition, and appearance. Simple fruits originate from a single set of seeds within one ovary, leading to uniform tissue layers derived from that ovary's pericarp and associated floral parts, whereas multiple fruits encompass seeds from multiple flowers, resulting in composite tissue layers from diverse ovaries and intervening receptacles.^[46] Externally, simple fruits exhibit a more straightforward morphology tied to a solitary flower's architecture, while multiple fruits display a complex, clustered form reflective of their inflorescence-based development. These distinctions have practical implications for botanical classification, as identification keys often require examining whether a fruit structure stems from a single flower or an inflorescence to differentiate simple from multiple types accurately.^[46]

With Aggregate Fruits

Aggregate fruits develop from a single flower that possesses multiple separate carpels, each maturing into a small fruitlet, resulting in a clustered structure.^[9] For instance, the raspberry consists of numerous drupelets, each derived from an individual carpel of one flower, forming an aggregate of drupes.^[47] Similarly, the strawberry is classified as an aggregate accessory fruit, where the true fruits are the achenes (each from a carpel) embedded in the enlarged receptacle tissue.^[48] The primary distinction between aggregate and multiple fruits lies in their floral origins: aggregate fruits arise from one flower with multiple ovaries, whereas multiple fruits form from an entire inflorescence of multiple flowers, each contributing one or more ovaries that fuse into a collective structure.^[49] This results in clustered drupes or similar units in aggregates versus a fused product from the clustered ovaries of many flowers in multiples.^[3] In multiple fruits, the anatomical fusion occurs across tissues from distinct flowers, contrasting with the single-flower origin in aggregates.^[46] Common misclassifications arise with fruits like the strawberry, which may appear as a multiple fruit due to its many visible achenes, but it is definitively an aggregate because all originate from the carpels of a single flower.^[45] Such confusion often stems from the accessory tissue's role in aggregates, leading to erroneous grouping with multiples that involve inflorescence fusion.^[50]

Biological and Ecological Importance

Dispersal Mechanisms

Multiple fruits exhibit diverse dispersal mechanisms adapted to their structural characteristics, primarily facilitating the spread of numerous seeds derived from clustered flowers. Animal-mediated dispersal is prevalent in fleshy multiple fruits, where the composite pericarp develops a sweet, colorful, and aromatic exterior to attract vertebrates such as birds and mammals. For instance, the pineapple (Ananas comosus), a multiple fruit formed from the fusion of berry-like structures, relies on animals consuming the juicy flesh, with viable seeds in wild varieties passing intact through digestive systems or being dropped nearby during feeding.^[51] Similarly, figs (Ficus spp.) and mulberries (Morus spp.), both multiple fruits composed of aggregated drupes, are ingested by frugivorous birds and bats, whose droppings deposit seeds over wide areas, enhancing germination rates due to scarification from gut passage.^[52] In some multiple fruits, irregular surfaces promote epizoochory, where fruits adhere to animal fur or skin.^[53] Abiotic dispersal mechanisms also occur in certain multiple fruits, particularly those with buoyant or lightweight structures. Water-mediated dispersal aids species in riparian or coastal habitats. For example, some fig species exploit hydrochory, where fruits float along rivers or during floods, allowing seeds to reach new germination sites distant from parent plants.^[54] These adaptations ensure broad dissemination, with the multiple fruit's buoyancy derived from air-trapping tissues in the fused receptacles.^[51] The composite structure of multiple fruits provides adaptive advantages for synchronized seed release, as the coordinated maturation of multiple ovaries into a single unit increases the overall attractiveness to dispersers and ensures mass seed output in one dispersal event. This aggregation amplifies the fruit's visibility and nutritional reward, promoting higher visitation rates by animals compared to isolated simple fruits, while the unified pericarp protects seeds during transport until release.^[51] In Osage orange, the large, multi-seeded synconium-like form historically facilitated long-distance dispersal by megafauna, though modern dispersal is limited.^[53] Overall, these mechanisms leverage the multiple fruit's morphology to optimize seed survival and colonization potential.

Evolutionary Significance

Multiple fruits evolved as a specialized reproductive strategy within angiosperms, originating from the fusion and maturation of ovaries across an entire inflorescence rather than a single flower. This developmental pathway links directly to inflorescence architecture and is evident in key families such as Moraceae (e.g., figs and mulberries) and Bromeliaceae (e.g., pineapples), where the trait likely diversified during the early Paleogene, around 50-60 million years ago, amid the radiation of tropical floras following the Cretaceous-Paleogene extinction.^[55]^[56] The adaptive advantages of multiple fruits include a substantial increase in seed count per fruiting structure, allowing plants to allocate resources efficiently for higher reproductive output in nutrient-limited tropical environments. By forming a composite mass, these fruits enhance dispersal efficiency through larger, more conspicuous packages that attract vertebrate dispersers, while the collective bulk provides mechanical protection against predation and environmental stressors for embedded seeds.^[57]^[58] Phylogenetically, multiple fruits are concentrated in tropical clades, particularly within the rosid order Rosales (Moraceae) and the commelinid order Poales (Bromeliaceae), reflecting convergent evolution in lineages adapted to megadiverse, animal-rich habitats. Fossil evidence, including syconium-like fruit imprints and dispersed achenes attributable to Moraceae from Eocene deposits in Europe and North America, corroborates their ancient origins and persistence in warm, humid paleoenvironments.^[56]^[59]

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