The family Cactaceae exhibits an assortment of fleshy and dry fruit types with various shapes dictated by the gynoecium outline and surrounding pericarpel. Consequently, conflicting terminology exists regarding cactus fruit classification because the fruit is a complex structure in which various floral parts participate in development. We examined fruit morphogenesis in four epiphytic cacti and provided a description of developmental events from post-anthesis to fruit maturation, which unveiled new structures valuable in fruit characterisation and taxonomy of the Hylocereeae and Rhipsalideae. Succinctly, the cactus fruit is a carpellar ovary embedded in a long-shoot (pericarpel). The pericarp originates from five components: internal ovarian epidermis that delimits the fruit locule, ovary (proper), collateral vascular bundles, pericarpel (receptacular origin), and external pericarpel epidermis. In addition, cell expansion and stored mucilage, a sticky substance involved in seed dispersal, occurs during fruit development. We propose the term cactidium, a complex fruit with accessory structures of pericarpellar origin surrounding the gynoecial boundary, to describe the cactus fruit. This term is appropriate because members of the Cactaceae bear unique traits, such as areoles in the reproductive structures (pericarpel), which may produce scale-leaves, bristles, and spines.
... The author distinguishes the simple fruits that originate from the superior ( Figure 1D) or semi-inferior ovary from those called pomaceous, which derive from the inferior ovary ( Figure 1E,F). Hitherto the various types of simple fruits have been studied from different families, such as Amaranthaceae [20,21], Asteraceae [22,23,24,25,26,27], Bignoniaceae [28,29,30], Cactaceae [31,32], Cordiaceae [33], Euphorbiaceae [34,35,36,37], Fabaceae [38,39,40], Lauraceae [41], Meliaceae [42], Moraceae [43], Myrtaceae [44], Nyctaginaceae [45,46]; Piperaceae [47,48], Rubiaceae [49] ( Figure 1E,F), and Santalaceae [50]. ...
... The former establishes that there is extensive fusion (connation and adnation) of the outer lower portions of the surrounding floral whorls to one another and to ovary wall (epigynous flower) ( Figure 1E); the latter assumes that carpels have phyletically "sunk" into tissue at the end of the cauline axis, with fusion of the receptacular tissue to the abaxial carpel wall [71]. Appendicular inferior ovaries may have occurred in ancestral Myrteae, Myrtaceae [44], and Cactaceae consists of a receptacular (pericarpel) inferior ovary [32]. ...
Angiosperm fruit originates from the ovary of the flower or it develops from the ovary, besides of the carpels, other inflorescence or floral parts may be involved in fruit formation; seed originates from the ovule. Four major classes of fruits are distinguished in angiosperms, viz. multiple fruits, aggregate fruits, schizocarps and simple fruits. Multiple fruits originate from inflorescences; the aggregate fruit comes from a single flower with apocarpous, pluricarpellate and pluripistillate gynoecium; schizocarps originate only from the ovary of a single flower, when ripe can separate into fragments called mericarps; and the simple fruits either are commonly originated from the ovary of a single unipistillate unicarpellate flower or syncarpic pluricarpellate flower. The diversity of seeds is also great among angiosperms, with bitegmic or unitegmic seeds, and alate, arillate, operculate, endothelial, pachychalazal, hairy or sarcotestal seeds. Reflections on fruit evolution are, in fact, very general, selecting a few examples that are restricted to families and genera. Fruit evolution is fundamentally based on changes of gynoecium types and differentiation of the pericarp. Among the hypotheses formulated in the literature, one predicts two main lines of evolution: in the first, there must have been a progressive reduction in the number of carpels that led to monocarpy, thus giving rise to the simple follicle; the second line must have led to a syncarpy, initially a loose adnation, which may have given rise to the capsule. The literature dealing with seed evolution in angiosperm taxa is very vast, but almost always not conclusive.
... The cactus ovary and fruit are exceptional structures surrounded by vegetative tissues, which in most cacti consist of nodes, internodes, axillary buds, and even rather ordinary leaves (Cota-Sánchez 2004). Recently, the fruit of the Cactaceae was defined as a cactidium, that is, a multifaceted carpellar ovary embedded in an elongated shoot (pericarpel) and the pericarp developing from five elements: internal ovarian epidermis, ovary (proper), collateral vascular bundles, pericarpel (of receptacular origin), and (Almeida et al. 2018). Other unique traits associated with the cactus flower and fruit are areoles, which may form scales, leaves, bristles, and spines. ...
... Thus, some anatomical fruit attributes vary throughout the family because the proportion of tissue(s) involved during development may be uneven. For instance, the fruits in tribes Rhipsalideae (Fig. 3e, g) and Hylocereeae ( Fig. 4e-h) share the uniseriate outer epidermis but are different in color, shape, and size because of the number of layers and presence/absence of the collenchymatous hypodermis (Almeida et al. 2018). ...
This review examines the historical research progress and areas of vivipary currently investigated in the Cactaceae. Vivipary, a rare attribute, has evolved multiple times in numerous plant lineages; however, a complete understanding of this event is still lacking in the cactus family and plants, in general.
This literature search combines the results obtained from scientific sources addressing aspects of vivipary published since 1900 to 2000, with an emphasis from 2000 to 2021. This systematic compendium summarizes findings in various aspects of vivipary, such as the taxonomic and ecological range, offspring survival, and the physiological bases of this phenomenon in the Cactaceae. To date, 77 viviparous taxa circumscribed in subfamilies Pereskioideae and Cactoideae are known, representing approximately 5.4% vivipary at the family level.
The taxonomic and geographic occurrence of this facultative reproductive attribute is discussed along with new reports, subsistence of viviparous and non-viviparous progeny, and a framework examining the phylogenetic distribution and putative origin of this generative mode in the family. The portrayal of the geographic distribution of viviparous species highlights the ubiquity of this trait and identifies vivipary hot spots in Cuba, the Brazilian Mata Atlântica, and NW Mexico, emphasizing ideas for cactus conservation. New data dealing with the role of the phytohormones abscisic acid and gibberellic acid in vivipary is examined in conjunction with the thermoregulatory properties of the fleshy viviparous fruits. Research areas deserving further studies are examined and several model species to conduct multidisciplinary research related to cactus vivipary in different areas of the Americas are proposed.
Sharable link at: https://rdcu.be/cVUvF
... The former element may be related to a smaller fruit with thinner pericarp and the latter to a larger incubation chamber for emerging seedling. Indeed, the development and resulting thickness of the cactus pericarp varies in different lineages of the family (Almeida et al. 2018). Thinner pericarp can facilitate seedling emergence as shown in several epiphytic cacti with thin, translucid fruit skin (Cota-Sánchez 2004), allowing germinated seeds inside the fruit to break through more easily as opposed to the thick, leathery pericarp of some viviparous cacti, such as Ferocactus herrerae J.G. Ortega ). ...
The basic aspects of vivipary, precocious germination within the fruit, are known. Consequently, research on this topic in the Cactaceae has increased in the last two decades and becoming more diversified. The family is amongst the most viviparous-rich angiosperm families together with some mangrove lineages. In this paper we report a new case of facultative vivipary, specifically cryptovivipary, in Cereus hildmannianus K. Schum., a South American columnar species and expand aspects regarding the physico-chemical traits of its fruits. The goals of this investigation were to: 1) report the first occurrence of vivipary in this species and characteristics of viviparous seedlings, and 2) describe some of the physical and chemical attributes of viviparous and nonviviparous fruits, such as size, weight, color, and total soluble solids (°Brix). Our findings show that this is the third account in Cereus Mill., for a 3% vivipary at the generic level. This discovery increases to 78 viviparous species for an overall 5.4% of viviparity family wide. Generally, the number and percentage of vivipary was low, with an average of 22.3 viviparous seedlings from an average of 1319 ungerminated seeds (= 1.7% vivipary/fruit). Statistical analyses indicate that non-viviparous fruits are larger, heavier, have higher content of soluble solids, thicker and brighter pericarp, and more seeds. Agriculturally, these attributes are more appealing to consumers suggesting that normal, non-viviparous fruits, are commercially more desirable; hence, vivipary is a detrimental character in fruit crops.
... The former element may be related to a smaller fruit with thinner pericarp and the latter to a larger incubation chamber for emerging seedling. Indeed, the development and resulting thickness of the cactus pericarp varies in different lineages of the family (Almeida et al. 2018). Thinner pericarp can facilitate seedling emergence as shown in several epiphytic cacti with thin, translucid fruit skin (Cota-Sánchez 2004), allowing germinated seeds inside the fruit to break through more easily as opposed to the thick, leathery pericarp of some viviparous cacti, such as Ferocactus herrerae J.G. Ortega ). ...
The basic aspects of vivipary, precocious germination within the fruit, are known. Consequently, research on this topic in the Cactaceae has increased in the last two decades and becoming more diversified. The family is amongst the most viviparous-rich angiosperm families together with some mangrove lineages. In this paper we report a new case of facultative vivipary, specifically cryptovivipary, in Cereus hildmannianus K. Schum., a South American columnar species and expand aspects regarding the physico-chemical traits of its fruits. The goals of this investigation were to: 1) report the first occurrence of vivipary in this species and characteristics of viviparous seedlings, and 2) describe some of the physical and chemical attributes of viviparous and nonviviparous fruits, such as size, weight, color, and total soluble solids (°Brix). Our findings show that this is the third account in Cereus Mill., for a 3% vivipary at the generic level. This discovery increases to 78 viviparous species for an overall 5.4% of viviparity family wide. Generally, the number and percentage of vivipary was low, with an average of 22.3 viviparous seedlings from an average of 1319 ungerminated seeds (= 1.7% vivipary/fruit). Statistical analyses indicate that non-viviparous fruits are larger, heavier, have higher content of soluble solids, thicker and brighter pericarp, and more seeds. Agriculturally, these attributes are more appealing to consumers suggesting that normal, non-viviparous fruits, are commercially more desirable; hence, vivipary is a detrimental character in fruit crops.
... The former element may be related to a smaller fruit with thinner pericarp and the latter to a larger incubation chamber for emerging seedling. Indeed, the development and resulting thickness of the cactus pericarp varies in different taxa(Almeida et al. 2018). Thinner pericarp can facilitate seedling emergence as shown in several epiphytic cacti with thin, translucid fruit skin (Cota-Sánchez 2004), allowing germinated seeds inside the fruit to break through more easily as opposed to the thick, leathery pericarp of some viviparous cacti, such as Ferocactus herrerae J.G. Ortega (Cota-Sánchez et al. 2007). ...
The basic aspects of vivipary, precocious germination within the fruit, are known. Consequently, research on this topic in the Cactaceae has increased in the last two decades and becoming more diversified. The family is amongst the most viviparous-rich angiosperm families together with some mangrove lineages. In this paper we report a new case of facultative vivipary, specifically cryptovivipary, in Cereus hildmannianus , a South American columnar species and expand aspects regarding the physico-chemical traits of its fruits. The goals of this investigation were to: 1) report the first occurrence of vivipary in this species and characteristics of viviparous seedlings, and 2) describe some of the physical and chemical attributes of viviparous and non-viviparous fruits, such as size, weight, color, and total soluble solids (°Brix). Our findings show that this is third account in Cereus , for a 3% vivipary at the generic level. This discovery increases to 78 viviparous species for an overall 5.4% of viviparity family wide. Generally, the number and percentage of vivipary was low, with an average of 22.3 viviparous seedlings from an average of 1319 ungerminated seeds (= 1.7% vivipary/fruit). Statistical analyses indicate that non-viviparous fruits are larger, heavier, have higher content of soluble solids, thicker and brighter pericarp, and more seeds. Agriculturally, these attributes are more appealing to consumers suggesting that normal, non-viviparous fruits, are commercially more desirable; hence, vivipary is a detrimental character in fruit crops.
... Fruits are showy with brilliant yellow or purple external exocarp with whitish or pink pulp ( Fig. 1) (Bauer 2003). The pulp in mature fruits is a combination of cells from the funiculus, endocarp and some of the innermost enlarged layers of the mesocarp that produced mucilage collapsed (Almeida et al. 2018). Based on a molecular phylogeny of tribe Hylocereeae, the delimitation of the closely related genera Hylocereus, Weberocereus and Selenicereus changed drastically (Korotkova et al. 2017). ...
Diverse tools and approaches are currently utilized to propose conservation strategies for ecosystems, areas and individual taxa. Here, ecological niche-based modeling, identification of areas of endemism, and diverse methods to determine conservation status are carried out to detect endangered species in Selenicereus. This genus in the Cactaceae has epiphytic species that are known for their edible fruit, called pitahayas or dragon fruit. With the exception of two species (S. grandiflorus and S. undatus), the other 21 studied species in Selenicereus were identified as threatened. Unique ecological niches were identified for these species, with implications for conservation. The most significant areas of species richness and endemism occur in Central America in unprotected areas, followed by other important regions in southern Mexico, which in contrast lie within reserves. Seasonal climates are characteristic of Selenicereus species commonly distributed in seasonally tropical dry forests and coastal vegetation, in areas in Central America where land transformation is rampant.
... та Phyllanthaceae Martinov (Gagliardi et al., 2014), Cactaceae Juss. (Almeida et al., 2018), Anacardiaceae R.Br. (Herrera et al., 2018), Calycanthaceae Lindl. ...
. In this review, the concept of fruit morphogenesis is treated in the context of implementation of the еvo-devo
approach in carpology. А new viewpoint on the fruit morphogenesis is proposed and justified, comprising the preanthetic, as well as post-anthetic periods of fruit development, id est, development of the gynoecium, and development
of the fruit itself. It is proposed to recognize ontogenetical (individual) and evolutionary (historical) aspects of fruit
morphogenesis, the first of them we can study directly, while the second aspect can be only hypothesized or treated as
a theoretical model of fruit evolution in consequence of some presumed changes in the individual fruit morphogenesis.
In this article these aspects are named as "ontomorphogenesis" and "phylomorphogenesis" of the fruit, correspondingly.
Our concept of ontomorphogenesis of the fruit involves four components that could not be brought together, such as
changes in the morphological structure of the gynoecium, abscission of the extragynecial floral parts and the style,
histogenesis of the fruit wall and other fruit parts, and terminal stages of the fruit morphogenesis (dehiscence, splitting,
or abscission). The current state of studies of these components in the individual and evolutionary contexts is discussed.
By examining the patterns of fruit evolution, we should consider factors acting at both the post-anthetic and pre-anthetic
periods of fruit ontomorphogenesis
Five Selenicereus species are well-known in the fruit market as dragon fruit, pitahaya, or pitaya. Native to the New World, pitahayas are considered underutilized crops with nutraceutical properties and easily propagated with a distribution that could potentially be extended to dry climates. Our goal is to understand the relationships of wild and cultivated populations and to determine genetic variation in a spatial scenario to discover hotspots of haplotype and genetic variation that will allow the conservation of valuable germplasm, as well as crop wild relatives. Sampling consisted of 170 individuals for three plastid molecular markers comprising the five cultivated species and as outgroups populations of four closely related species were included in the haplotype analyses. Genealogical relationships were determined, along with genetic variation in spatial patterns. The majority of the haplotypes were shared among the nine species in a geographic pattern; however, distant populations of different species also shared haplotypes. Selenicereus monacanthus displayed the highest genetic variation; its haplotype network is complex and intricate, probably related to the management to which the populations have been subjected, in which certain attributes suitable for cultivation and valuable for the fruit market have been selected. Historical evidence suggests that S. undatus has been cultivated in home gardens in the Maya area since pre-Columbian times, and the highest genetic diversity was found there. Conservation of wild crop relatives is important to preserve underutilized crops, therefore southern Mexico and northern Central America are the most relevant regions to protect genetic diversity of pitahayas.
Key message
Contrasting morphologies in Disocactus are the result of differential development of the vegetative and floral tissue where intercalary growth is involved, resulting in a complex structure, the floral axis.
Abstract
Species from the Cactaceae bear adaptations related with their growth in environments under hydric stress. These adaptations have translated into the reduction and modification of various structures such as leaves, stems, lateral branches, roots and the structuring of flowers in a so-called flower-shoot. While cacti flowers and fruits have a consistent structure with showy hermaphrodite or unisexual flowers that produce a fruit called cactidium, the developmental dynamics of vegetative and reproductive tissues comprising the reproductive unit have only been inferred through the analysis of pre-anthetic buds. Here we present a comparative analysis of two developmental series covering the early stages of flower formation and organ differentiation in Disocactus speciosus and Disocactus eichlamii, which have contrasting floral morphologies. We observe that within the areole, a shoot apical meristem commences to grow upward, producing lateral leaves with a spiral arrangement, rapidly transitioning to a floral meristem. The floral meristem produces tepal primordia and a staminal ring meristem from which numerous or few stamens develop in a centrifugal manner in D. speciosus and D. eichlamii, respectively. Also, the inferior ovary derives from the floral meristem flattening and an upward growth of the surrounding tissue of the underlying stem, producing the pericarpel. This structure is novel to cacti and lacks a clear anatomical delimitation with the carpel wall. Here, we present a first study that documents the early processes taking place during initial meristem determination related to pericarpel development and early floral organ formation in cacti until the establishment of mature floral organs.
The genus Opuntia, commonly known as prickly pear cactus, includes species that produce nutritious fruits and young, edible cladodes (stem pads, also called joints), which are used as a vegetable. The prickly pear fruit is known as tuna, Indian fig, Christian fig, and tuna de Castilla. Mexico is considered one of the major areas of genetic diversity of Opuntia, and Opuntia ficus-indica is one of the most agro-economically important cactus crop species and is cultivated in arid and semiarid regions of the world for its fruits. The spineless forms correspond to horticultural varieties. The prickly pear fruit is divided into three components that may be exploited commercially: seeds, peel, and pulp. This fruit contains approximately 85% water, 15% sugar, 0.3% ash, and less than 1% protein. The flesh is a good source of minerals and several types of amino acids (alanine, arginine, and asparagine). Important vitamins include vitamin C (ascorbic acid), E, K, and beta-carotenes. Flavonoids, effective antioxidants, are another important constituent. The betalain pigments are responsible for the colors of the fruit and also have antioxidant properties. The general distribution of nutrients and antioxidants in the fruit is an indication that the ingestion of the whole fruit is more beneficial from a health perspective because more potentially nutraceutical active ingredients are absorbed and used by our bodies. Considering the chemical components of the prickly pear cactus, its nutritional capacity is relatively modest and should be used as a dietary complement. In view of the popular and increasing trend in the demand for nutraceuticals and increased desire for natural ingredients and food products promoting health, the multiple functional properties of cactus pear in conjunction with its antioxidant properties fit well with this trend. Furthermore, the prickly pear fruit can be considered as "the bridge of life" because it is the only food and water resource for animals during the long dry seasons in the deserts.
Pereskia diaz-romeroana and its putative relatives are scandent plants with vigorously growing long shoots, small flowers and fruits, and tuberous root systems. Except for size, floral development parallels that in other pereskias. The gynoecium has three to six carpels, is basically superior with axlie placentation, and is partially divided into locules by the connate carpel margins. The closest relative of these Andean forms appears to be P. aculeata.
Boke, Norman H. (U. Oklahoma, Norman.) Anatomy and development of the flower and fruit of Pereskia pititache. Amer. Jour. Bot. 50 (8): 843–858. Illus. 1963.—Flowers of P. pititache are about 6 cm in diameter and perigynous. The receptacle bears numerous broad bracts; the inner perianth segments are orange and deeply cleft; the numerous stamens develop centrifugally. The fundamentally superior gynoecium is broad and flat and consists of 10–18 connate carpels, the fertile portions of which are involute and adnate to the conical floral axis. The 8–16 ovules in each of the pocket-like locules are borne in 2 rows along the zone of adnation; placentation is axile. The short style and the style branches are lined with stigmatic tissue, which extends downward among the ovules. There is no definite stigma. The tip of the floral axis retains its meristematic characteristics beyond anthesis. In early fruit development, the rim of the floral cup grows in height, while the residual floral axis becomes a conspicuous peg-like columella. Concomitantly, the formation of numerous mucilage cells and cavities causes the ovary partitions and parts of the ovary roof to disintegrate. As a result, the seeds are contained in a single, moat-like cavity, which appears inferior. Late fruit development is characterized by the differentiation of numerous fiber sclereids in association with the extensive and complex vascular system.
Studies on breeding systems and flower morphology are valuable to infer how environmental factors impose evolutionary change in plants. This study focused on the characterization of floral morphs and reproductive systems in Pachycereus pringlei and how this iconic columnar cactus might be a useful genus to understand the evolution of these highly variable structures. We determined breeding systems, characterized floral morphs in five genetic populations, and used pollen:ovule ratios and stamen-stigma distance to verify the sexual system in bisexual flowers. Our inquiries provide insights into the factors driving intra-specific disparity in flower attributes and the reproductive versatility in P. pringlei. Foremost, the lability of breeding systems is expressed primarily as gynodioecy in the North, trioecy in the South, mainly dioecy in CBS, and hermaphroditic in Catalana and Cerralvo Islands. A latitudinal trend in ovule production, dimensions in gynoecium, androecium, and floral display characters is consistent with a northward increase in vegetative traits suggesting physiological responses to the environmental variation characterizing the Baja California Peninsula (BCP). Also, nectary size decreases northwards in staminate flower, but these flowers are larger in the CBS and South populations suggesting sex-specific selection, e.g., pollinator-driven, acting in different magnitudes on sexual attributes of floral morphs and populations. The presence of rudimentary structures of the dysfunctional sex support the hypothesis of the evolution of unisexuality from an early bisexual ancestor. In conclusion, this investigation provides insights into the factors driving intra-specific disparity in flower attributes and the reproductive versatility in P. pringlei to replace ancestral conditions, specifically the substitution of the hermaphrodite phase with dioecious, gynodioecious, and trioecious breeding systems throughout mainland BCP and Sonora. We posit that the biogeographic patterns of breeding systems and floral traits of this emblematic cactus resulted from the interaction of past factors (northwards range expansion) and contemporary biotic and abiotic factors.
Cactaceae is classified within the order of Caryophyllales and includes approximately 1,600 species of plants. Phylogenetic placement within the Caryophyllales is undisputed, because cacti and other families within the order share synapomorphies or derived characters. An example is the occurrence of betalains, a class of nitrogenous pigments derived from tyrosine. This chapter discusses the evolution of the Cactaceae family, paying special attention to molecular and genetic approaches for analyzing the morphological distinctiveness and monophyly of Cactaceae. It highlights the importance of molecular systematic studies in providing new insights into the origin and diversification of cacti. The combination of molecular, morphological, and biogeographical data provide the best estimate of phylogeny within the Cactaceae and a more reliable source of biological information about this family.
![General flower and fruit morphology of Epiphyllum phyllanthus (A–E), Hylocereus undatus (F–I), Lepismium warmingianum (J–M), and Rhipsalis cereuscula (N–Q). (A) Flower in lateral view. (B) Detail of perianth. (C) Young fruit. (D) Mature fruit. (E) Longitudinal section of a mature fruit. (F) Flower. (G) Immature fruit. (H) Mature fruit on phylloclade. (I) Lateral and longitudinal view of a mature fruit. (J) Phylloclade with flower and floral buds. (K) Flower close-up. (L) Post-anthesis and young fruit. (M) Immature and mature fruits. (N) General view of the plant. (O) Close-up of a flower. (P) Post-anthesis and young fruit. (Q) Mature fruit. Scale bars = 4 cm (A, F, G, I); 1.5 cm (B); 1 cm (C, K, M); 2 cm (D, E, J): 5 cm (H, N); 0.5 cm (L, O–Q). [Colour online.]](https://www.researchgate.net/profile/J-Hugo-Cota-Sanchez/publication/325730401/figure/fig1/AS:11431281290928983@1731873267988/General-flower-and-fruit-morphology-of-Epiphyllum-phyllanthus-A-E-Hylocereus-undatus_Q320.jpg)
![Fruit anatomy of Epiphyllum phyllanthus in cross-section. (A and B) Young fruit, post-anthesis, general view (A) and detail of pericarp wall (B). (C and D) General view of immature fruit and detail of pericarp wall. (E) Mature fruit and detail of pericarp wall; ex, exocarp; pc, pericarp; pe, pericarpel origin; ov, ovarian origin; sc, mucilage secretory cavity. Scale bars = 1 mm (A–D); 1.8 mm (E). [Colour online.]](https://www.researchgate.net/profile/J-Hugo-Cota-Sanchez/publication/325730401/figure/fig2/AS:11431281290890787@1731873269367/Fruit-anatomy-of-Epiphyllum-phyllanthus-in-cross-section-A-and-B-Young-fruit_Q320.jpg)
![Fruit anatomy of Hylocereus undatus in longitudinal (A, B, E, and I–L) and cross-section (C, D, F, G, and H). (A–H) Young fruit (post-anthesis). (B) Detail of the outermost parenchyma layers of pericarp. (C) Detail of the outermost region of young pericarp (outlined in Fig. 3H). (D) Innermost part of pericarp and funiculus. (E) Detail of epidermis and collenchymatous hypodermis. (F) Detail of a large vascular bundle between the outer and inner layers of the pericarp (outlined in Fig. 3H). (G) Detail of funiculus cells and a vascular bundle. (H) Young fruit (post-anthesis), longitudinal section. (I–L) Mature fruit. (J) Detail of the outer part of the pericarp. (K) Detail of the middle part of the pericarp. (L) Detail of the inner part of the pericarp; cu, cuticle; ex, exocarp; fu, funiculus; hp, hypodermis; ov, ovarian origin; pc, pericarp; pe, pericarpel origin; sl, scale-leaf; vb, vascular bundle. Scale bars = 2 mm (A, H, I); 100 μm (B); 400 μm (C, D, F); 50 μm (E); 200 μm (G, J–L). [Colour online.]](https://www.researchgate.net/profile/J-Hugo-Cota-Sanchez/publication/325730401/figure/fig3/AS:11431281290939419@1731873271414/Fruit-anatomy-of-Hylocereus-undatus-in-longitudinal-A-B-E-and-I-L-and-cross-section_Q320.jpg)
![Structural organization of Lepismium warmingianum fruit in longitudinal (A, E, and I) and cross-sections (B–D, F–H, and J–L) and SEM view (D, H, and L). (A and B) Post-anthesis young fruit. (C) Detail of pericarp (pericarpel + ovary). (D) Pericarp and ovules. (E and F) Young fruit. (G) Detail of pericarp. (H) Ovules within young fruit. (I and J) Mature fruit. (K) Detail of pericarp. (L) Pericarp and seed of mature fruit; en, endocarp; ex, exocarp; ov, ovarian origin; pc, pericarp; pe, pericarpel origin; sc, mucilage secretory cavity. Scale bars = 2 mm (A, E, I, J); 1 mm (B, F); 200 μm (C); 300 μm (G, K); 250 μm (D); 500 μm (H, L). [Colour online.]](https://www.researchgate.net/profile/J-Hugo-Cota-Sanchez/publication/325730401/figure/fig4/AS:11431281290910319@1731873273467/Structural-organization-of-Lepismium-warmingianum-fruit-in-longitudinal-A-E-and-I-and_Q320.jpg)
![Structural organization of Rhipsalis cereuscula fruit in longitudinal (A, E, I) and cross (B–D, F–H, J–L) sections and SEM view (D, H, L). (A and B) Post-anthesis young fruit. (C) Detail of pericarp. (D) Detail of pericarp and ovules. (E and F) Young fruit. (G) Detail of pericarp. (H) Wall and ovules of young fruit. (I and J) Mature fruit. (K) Detail of pericarp. (L) Pericarp and seeds surrounded by mucilage in mature fruit. en, endocarp; ex, exocarp; mu, seed mucilaginous sheath; ov, ovarian origin; pc, pericarp; pe, pericarpel origin; sc, secretory cavity; rv, recurrent vascular bundle. Scale bars = 1 mm (A, B, F); 300 μm (C); 1.5 mm (E, I, J); 200 μm (G, K); 400 μm (H); 500 μm (D, L). [Colour online.]](https://www.researchgate.net/profile/J-Hugo-Cota-Sanchez/publication/325730401/figure/fig5/AS:11431281290881085@1731873274511/Structural-organization-of-Rhipsalis-cereuscula-fruit-in-longitudinal-A-E-I-and-cross_Q320.jpg)
