Pericarp histogenesis and histochemistry during fruit development in Butia capitata (Arecaceae)

Palm fruits show great structural complexity, and in-depth studies of their development are still scarce. This work aimed to define the developmental stages of the fruit of the neotropical palm Butia capitata and to characterize the ontogenesis of its pericarp. Biometric, anatomical, and histochemical evaluations were performed on pistillate flowers and developing fruits. The whole fruit develops in three phases: (I) histogenesis (up to 42 days after anthesis – DAA), when the topographic regions of the pericarp are defined; (II) pyrene maturation (42 to 70 DAA), when the sclerified zone of the pericarp is established; and (III) mesocarp maturation (70 to 84 DAA), when reserve deposition is completed. During pericarp ontogenesis (i) the outer epidermis and the outer mesophyll of the ovary give origin to the exocarp (secretory epidermis, collenchyma, parenchyma, sclerenchyma, and vascular bundles); (ii) the median ovarian mesophyll develops into the mesocarp, with two distinct topographical regions; (iii) the inner ovarian epidermis originates the endocarp; and in the micropylar region, it differentiates into the germination pore plate, a structure that protects the embryo and controls germination. (iv) Most of the inner region of the mesocarp fuses with the endocarp and, both lignified, give rise to the stony pyrene; (v) in the other regions of the mesocarp, carbohydrates and lipids are accumulated in a parenchyma permeated with fiber and vascular bundles. The development of the B. capitata pericarp presents high complexity and a pattern not yet reported for Arecaceae, which supports the adoption of the Butia-type pyrenarium fruit class.

     

Development of ovule,fruit and seed of Xyris (Xyridaceae,Poales) and taxonomic considerations

The development of the ovule, fruit and seed of Xyris spp. was studied to assess the embryological characteristics of potential taxonomic usefulness. All of the studied species have (1) orthotropous, bitegmic and tenuinucellate ovules, with a micropyle formed by both the endostoma and exostoma; (2) a cuticle in the ovules and seeds between the nucellus/endosperm and the inner integument and between the inner and outer integuments; (3) helobial, starchy endosperm; (4) a reduced, campanulate and undifferentiated embryo; (5) a seed coat formed by a tanniferous endotegmen, endotesta with thick‐walled cells and exotesta with thin‐walled cells; and (6) a micropylar operculum formed from inner and outer integuments. The pericarp is composed of a mesocarp with cells containing starch grains and an endocarp and exocarp formed by cells with U‐shaped thickened walls. The studied species differ in the embryo sac development, which can be of the Polygonum or Allium type, and in the pericarp, which can have larger cells in either endocarp or exocarp. The Allium‐type embryo sac development was observed only in Xyris spp. within Xyridaceae. Xyris also differs from the other genera of Xyridaceae by the presence of orthotropous ovules and a seed coat formed by endotegmen, endotesta and exotesta, in agreement with the division of the family into Xyridoideae and Abolbodoideae. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 619–628.     

Fruit micromorphology and anatomy of Valeriana officinalis s. str. (Valerianaceae)

Fruits of two varieties of Valeriana officinalis s. str. (var. officinalis , var. nitida ) are similar in general construction, but differ in details of external and internal structure. The outer cells of the pericarp form a regularly punctuated surface in both taxa. Scanning electron microscopy demonstrates variation in cuticular sculpturing of the outer epidermal cell walls and the presence of epicuticular wax. The surface of fruit hairs varies from micropapillate in var. officinalis to linear warty in var. nitida . In the mature rericarp there occur three distinct histological zones: an outer exocarp, a central mesocarp, and an inner endocarp. The seed is small, enclosed in the indehiscent fruit, with thin seed coat and a straight embryo. Endosperm is absent. The results of this carpological study, especially the SEM characters of pericarp surface, may provide criteria useful for delimitation of V officinalis varieties.     

Pericarp development and fruit structure in borassoid palms (Arecaceae-Coryphoideae-Borasseae)

Background and Aims

The Borasseae form a highly supported monophyletic clade in the Arecaceae–Coryphoideae. The fruits of Coryphoideae are small, drupaceous with specialized anatomical structure of the pericarp and berries. The large fruits of borassoid palms contain massive pyrenes, which develop from the middle zone of the mesocarp. The pericarp structure and mode of its development in Borasseae are similar to those of Eugeissona and Nypa. A developmental carpological study of borassoid palms will allow us to describe the process of pericarp development and reveal the diagnostic fruit features of borassoid palms, determine the morphogenetic fruit type in Borasseae genera, and describe similarities in fruit structure and pericarp development with other groups of palms.

Methods

The pericarp anatomy was studied during development with light microscopy based on the anatomical sections of fruits of all eight Borasseae genera.

Key Results

The following general features of pericarp structure in Borasseae were revealed: (1) differentiation of the pericarp starts at early developmental stages; (2) the exocarp is represented by a specialized epidermis; (3) the mesocarp is extremely multilayered and is differentiated into several topographical zones – a peripheral parenchymatous zone(s) with scattered sclerenchymatous elements and vascular bundles, a middle zone (the stony pyrene comprising networks of elongated sclereids and vascular bundles) and an inner parenchymatous zone(s); (4) differentiation and growth of the pyrene tissue starts at early developmental stages and ends long before maturation of the seed; (5) the inner parenchymatous zone(s) of the mesocarp is dramatically compressed by the mature seed; (6) the endocarp (unspecialized epidermis) is not involved in pyrene formation; and (7) the spermoderm is multilayered in Hyphaeninae and obliterated in Lataniinae.

Conclusions

The fruits of Borasseae are pyrenaria of Latania-type. This type of pericarp differentiation is also found only in Eugeissona and Nypa. The fruits of other Coryphoideae dramatically differ from Borasseae by the pericarp anatomical structure and the mode of its development.     

Pericarp formation in early divergent species of Arecaceae (Calamoideae,Mauritiinae) and its ecological and phylogenetic importance

Lepidocaryum tenue, Mauritia flexuosa and Mauritiella armata belong to the subtribe Mauritiinae, one early divergent lineage of the Arecaceae and one of the few of Calamoideae that occur in South America. These species occur in swampy environments and have fruits that are characteristically covered with scales. The objective of this study was to describe the formation of the layers of the pericarp within this subtribe and attempt to correlate fruit structure with the environment where species typically occur. Toward this goal, flowers in pre-anthesis and anthesis and fruits throughout development were analyzed using standard methods for light microscopy. The ontogeny of the layers of the pericarp of all three species was found to be similar. The scales were formed from non-vascularized emergences composed of exocarp and mesocarp. The median mesocarp accumulates lipids only in M. flexuosa and M. armata. The inner mesocarp together with the endocarp becomes papyraceous and tenuous in all species. This internal region of pericarp showed collapsed cells due to seed growth at the end of fruit development. Fruits of Mauritiinae are baccate, and the characters of the pericarp, especially the inner mesocarp and endocarp, help to maintain moisture. On the other hand, many species close to Mauritiinae show pericarp with sclerenchyma adjacent to the seed. This variation can contribute to understand the importance of this striking character in dispersal, germination and colonization in Arecaceae.     

龙眼果皮形态结构比较观察及其与果实耐贮运的关系

比较了福建省 1 0个主栽龙眼品种果实的果皮形态和结构 ,结果表明 :不同品种在果皮厚度、外果皮表面颜色、龟状纹、放射线、瘤状突、刺毛、外果皮皮孔、周皮层厚度、栓质层厚度和连续性、中果皮薄壁组织细胞排列、石细胞大小、含量、排列和分布 ,维管束发达状况、排列和分布 ,内果皮表皮细胞排列和角蜡质层厚度等方面均存在着明显差异。风梨味、东壁、油潭本、乌龙岭、红核子、蕉眼龙眼果皮厚 ,外果皮表面瘤状突和剌毛多 ,外果皮周皮层、栓质层厚且连续性好 ,中果皮石细胞 (团 )含量多且排列紧密 ,分布在中果皮外侧且在中果皮中所占比例大 ,维管束发达且排列有序 ,内果皮角蜡质层厚 ;这些品种果实耐贮运、抗病性强。而水涨、赤壳、福眼、普明庵龙眼果皮薄 ,外果皮周皮层薄、栓质层不发达 ,中果皮石细胞 (团 )含量少、分布分散 ,维管束不发达 ,薄壁组织细胞胞间隙大 ,皮孔间隙大、皮孔通道与中果皮组织细胞间隙相通 ;这些品种的果实不耐贮运、抗病性弱。讨论了龙眼外果皮表面主色为褐色和内果皮比外果皮更容易褐变的解剖学原因及龙眼果皮形态结构与果实耐贮运的关系。     

Complex combination of seed dormancy and seedling development determine emergence of Viburnum tinus (Caprifoliaceae)

BACKGROUND AND AIMS: The shrub Viburnum tinus is widely distributed in mattoral vegetation of the Mediterranean basin. The purpose of the present study was to classify the seed dormancy type and examine the requirements for embryo growth, root protrusion and shoot emergence. METHODS: Overwintered fruits were collected in western Spain in April 2001 and prepared in three ways: entire pericarp was removed, exocarp and mesocarp were removed or fruits were left intact. Fruits treated in these three ways were subjected to artificial annual temperature cycles or to constant temperature regimes for 1.5 years. KEY RESULTS: Removal of exocarp and mesocarp was necessary for embryo growth and germination. High temperature favoured dormancy alleviation and embryo growth, intermediate to low temperatures favoured root protrusion, and intermediate temperature shoot emergence. There was substantial germination at constant temperature regimes, indicating an overlap between temperature intervals suitable for the different stages of embryo and seedling development. Functionally, V. tinus has the same root and shoot emergence pattern that is described for other Viburnum species considered to have epicotyl dormancy. However, the requirement for high and low temperatures for radicle protrusion and epicotyl emergence, respectively, was missing in V. tinus; these characters are the foundation for the epicotyl dormancy classification. CONCLUSIONS: It is concluded that V. tinus does not have epicotyl dormancy. Instead, there is a combination of a weak morphophysiological dormancy and a slow germination process, where different temperatures during an annual cycle favour different development stages. The present study suggests that the first complete seedlings would emerge in the field 1.5 years after fruit maturation in October, i.e. seed dispersal during winter, embryo growth during the first summer, root protrusion and establishment during the second autumn and winter, and cotyledon emergence during the second spring.     

Pericarp anatomy of wild roses (Rosa L., Rosaceae)

The pericarp anatomy of representatives of all subgenera and sections of the genus Rosa was studied. All species have the same basic pericarp structure: it is composed of inner and outer endocarps, mesocarp and exocarp formed by the epidermis and hypodermis. The differences concern mainly the thickness of particular layers, and the shape and size of their cells. Cells of the endocarp and mesocarp are thick-walled. The only exception is Rosa rugosa mesocarp, which is composed of rather thin-walled cells with a large lumen. The endocarp structure of Rosa achenes resembles the drupe of the genus Prunus s.l. and drupelets of Rubus species.     

Ovule,fruit and seed development in Abolboda (Xyridaceae,Poales): implications for taxonomy and phylogeny

Xyridaceae belongs to the xyrid clade of Poales, but the phylogenetic position of the xyrid families is only weakly supported. Xyridaceae is divided into two subfamilies and five genera, the relationships of which remain unclear. The development of the ovule, fruit and seed of Abolboda spp. was studied to identify characteristics of taxonomic and phylogenetic value. All of the studied species share anatropous, tenuinucellate and bitegmic ovules with a micropyle formed by the inner and outer integuments, megagametophyte development of the Polygonum type, seeds with a tanniferous hypostase, a helobial and starchy endosperm and an undifferentiated embryo, seed coat derived from both integuments with a tanniferous tegmen and a micropylar operculum, and fruits with a parenchymatous endocarp and mesocarp and a sclerenchymatous exocarp. Most of the ovule and seed characteristics described for Abolboda are also present in Xyris and may represent a pattern for the family. Abolboda is distinguished by the ovule type, endosperm formation and the number of layers in the seed coat, in agreement with its classification in Abolbodoideae. The following characteristics link Xyridaceae to Eriocaulaceae and Mayacaceae, supporting the xyrid clade: tenuinucellate, bitegmic ovules; seeds with a tanniferous hypostase, a starchy endosperm and an undifferentiated embryo; and a seed coat with a tanniferous tegmen. A micropylar operculum in the seeds of Abolboda is described for the first time here and may represent a synapomorphy for the xyrids. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 144–154.     

Genetic separation of autonomous endosperm formation (AutE) from the two other components of apomixis in Hieracium

In apomictic Hieracium subgenus Pilosella species, embryo sacs develop in ovules without meiosis. Embryo and endosperm formation then occur without fertilization, producing seeds with a maternal genotype encased in a fruit (achene). Genetic analyses in H. praealtum indicate a dominant locus (LOA) controls meiotic avoidance, and another dominant locus (LOP) controls both fertilization-independent embryogenesis and endosperm formation. While cytologically examining developmental events in ovules of progeny from crosses between different wild-type and mutant Hieracium apomicts, and a sexual Hieracium species, we identified two plants, AutE196 and AutE24, which have lost the capacity for meiotic avoidance and fertilization-independent embryo formation. AutE196 and AutE24 exhibit autonomous endosperm formation and set parthenocarpic, seedless achenes at a penetrance of 18 %. Viable seed form after pollination. Cytological examination of 102 progeny from a backcross of AutE196 with sexual H. pilosella showed that autonomous endosperm formation is a heritable, dominant, qualitative trait, detected in 51 % of progeny. Variation in quantitative trait penetrance indicates other factors influence its expression. The correlation between autonomous endosperm development and mature parthenocarpic achene formation suggests the former is sufficient to trigger fruit maturation in Hieracium. The developmental component of autonomous endosperm formation is therefore genetically separable from those controlling meiotic avoidance and autonomous embryogenesis in Hieracium and has been denoted as AutE. We postulate that tight linkage of AutE and genes controlling autonomous embryogenesis at the LOP locus in H. praealtum may explain why inheritance of autonomous seed formation is typically observed as a single component.     

Accrescimento del pericarpo,seme, endosperma ed embrione in prunus Amygdalus stokes

Abstract

GROWTH OF PERICARP, SEED, ENDOSPERM, AND EMBRYO IN PRUNUS AMYGDALUS STOKES. — The fruits of an almond-tree growing at Bari were collected weekly from February 22nd to July 11th and on August 16th 1960. The material was kept in fixative; the growth of the various organs was studied both from a morphological and a quantitative point of view. Special attention was given to growth of the endosperm, especially during the nuclear stage and at the beginning of cellularisation (Figg. 1-14), and to the developement of the embryo until it reaches the « heart-shaped » stage (Figg. 15–22). From a quantitative point of view, the volume and main diameters of pericarp and seed, and whenever possible endosperm and embryo, were measured for each fruit. Most of the data are given in Tables I to V and Figg. 23 and 24.

If reference is made to the 3 phases of fruit growth established for other species (notably peaches and cherries), the main conclusions are that:

1. phase I (growth of pericarp, testa and nucellus) is clearly recognisable; it ends after the micropylar portion of the endosperm has become cellular and the embryo heart shaped;

2. phase II is also present: during this phase most of the growth of endosperm and embryo takes place; while the seed has reached its definite size at the end of phase I, the pericarp undergoes a period of greatly reduced growth;

3. two weeks after the beginning of phase II the pericarp seems to resume growth just for a very short period, judging at least by the weekly values of the ratio pericarp volume to seed volume (see Fig. 23); this seems to indicate the existence of a new phase, that is phase III, which in fleshy fruits of the genus Prunus corresponds to a much longer and important process of pericarp growth than in the almond;

4. as in the peaches and cherries therefore a crisis in pericarp growth occurs during the period of maximum rate of growth of the cellular endosperm and embryo;

5. the sequence: cellularisation of the endosperm, growth of endosperm and embryo, ceasing of seed growth, and reduction in pericarp growth is very clear, particularly if we take into account growth in length rather than in volume; both morphological and quantitative data would indicate the importance of the endosperm not only for the beginning of embryo development, but also for the control of pericarp growth.

     

Gibberellin-like Activity in the Developing Avocado Fruit

Gibberellin-like activity in avocado (Persea americana) fruit extracts was measured by the barley endosperm bioassay. Fruit tissues were anlyzed separately during fruit growth. The level of the activity found was very high in the endosperm and in the seed coats, but decreased in the latter during fruit growth. No measurable gibberellin-like activity was detected in the mesocarp or in the embryo. It is assumed that the seed coats are a site of production of gibberellin-like substances in the avocado fruit.     

Structure of the unusual explosive fruits of the early diverging angiosperm Illicium (Schisandraceae s.l., Austrobaileyales)

All Illicium spp. have explosive fruits, which is a unique character among the basal grade of angiosperms. Illicium fruits consist of several ventrally dehiscing follicles developing from conduplicate carpels, with a prominent, slightly postgenitally fused ventral slit. The closure of the ventral slit is also secured by two mirror‐symmetrical massive longitudinal sclerenchymatous bands in the mesocarp along the edges and by turgor pressure. The pericarp differentiates into a fleshy (or coriaceous) peripheral zone (exocarp and mesocarp) with numerous ethereal‐oil‐containing cells and a sclerenchymatous (single‐layered, palisade) inner zone (endocarp). Dehydration of the fleshy zone of the pericarp and partial compression of the epidermal sclereids with U‐shaped wall thickenings lining the ventral suture are instrumental in explosive fruitlet dehiscence. Generally, the fruit structure of Illicium differs dramatically from those in other early diverging angiosperms. Gynoecium and fruit structure (and a probable early Cretaceous divergence from the Schisandra‐Kadsura clade) provide evidence for treatment of Illicium as separate from Schisandraceae s.s. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013 , 171 , 640–654.     

Development and structure of the pericarp of Lannea discolor (Sonder) Engl.(Anacardiaceae)

Development and structure of the pericarp of Lannea discolor (Sonder) Engl.(Anacardiaceae). The exocarp develops from the outer epidermis and subepidermal, parenchymatous cell layers of the ovary wall. A parenchymatous zone with secretory cavities more or less delimits the exocarp internally. The inner part of the parenchymatous mesocarp is tanniniferous. The parenchymatous transition zone between mesocarp and sclercnchymatous endocarp or sderocarp, contains vascular tissue. The inner endocarp and operculum develop from the inner epidermis and subepidermal parenchyma of the ovary wall, while the outer endocarp develops from the parenchymatous zone with procambium strandS. Comparing the pericarp of L.discolor with those of Sclerocarya birrea subsp. caffra and Rhus lancea , the close affinity with Sclerocarya birrea subsp. caffra is evident.     

Significance of achene characteristics and within-achene resource allocation in the germination strategy of tetraploid Aster pilosus var. pilosus (Asteraceae)

The interrelationships among achene weight, allocation to embryo and pericarp, and germination time were determined for 500 stratified achenes of tetraploid Aster pilosus Willd. var. pilosus. Only 52.6% of the achenes germinated. Germinated achenes were significantly heavier than ungerminated achenes. Germination time was independent of achene weight and embryo weight, but varied inversely with pericarp weight. Variable achene weight is evolutionarily advantageous. Heavy achenes are at an advantage in that their proportionately larger embryos and thinner pericarps facilitate germination, promoting competitive establishment of seedlings. Lighter achenes are also at an advantage through increased dispersibility, and their relatively thick pericarp provides a persistent seed bank. Evolutionary pressures presumably maintain the variability in achene weight of var. pilosus. These results are discussed in the context of the early midsuccessional ecology of var. pilosus.     

Development and structure of the drupe in Sclerocarya birrea (Richard) Hochst. subsp. caffra Kokwaro (Anacardiaceae), with special reference to the pericarp and the operculum

The development and structure of the exo-, meso- and endocarp of the drupe of Sclerocarya birrea subsp. caffra were examined. The mature exocarp comprises the outer epidermis with stomata and lenticels, subepidermal collenchyma and parenchymatous layers with secretory canals. This exocarp sensu lato develops from the outer epidermis and the outer layers of the ovary wall. The fleshy parenchymatous mesocarp or sarcocarp also contains secretory tissue. The mesocarp develops after endocarp differentiation and lignification. The developmental sequence within the pericarp corresponds to the general pattern in drupes. The endocarp or sclerocarp, which is not stratified, consisting mainly of brachysclereids, fibres and vascular elements, develops from the inner epidermis and adjacent tissue of the young ovary wall including the procambium strands. The operculum represents a well-defined part of the endocarp. Early in its development a parenchymatous zone already clearly demarcates the operculum. The literature on the pericarp of the Anacardiaceae drupe is discussed to establish the diagnostic value of these morphological characteristics for future taxonomic studies.     

Fruit structure of Amborella trichopoda (Amborellaceae)

The indehiscent fruitlets of the apparently basalmost extant angiosperm, Amborella trichopoda, have a pericarp that is differentiated into five zones, a thin one‐cell‐layered skin (exocarp), a thick fleshy zone of 25–35 cell layers (outer mesocarp), a thick, large‐celled sclerenchymatous zone (unlignified) of 6–18 cell layers (middle mesocarp), a single cell layer with thin‐walled (silicified?) cells (inner mesocarp), and a 2–4‐cell‐layered, small‐celled sclerenchymatous zone (unlignified) derived from the inner epidermis (endocarp). The border between inner and outer mesocarp is not even but the inner mesocarp forms a network of ridges and pits; the ridges support the vascular bundles, which are situated in the outer mesocarp. In accordance with previous observations by Bailey & Swamy, no ethereal oil cells were observed in the pericarp; however, lysigenous cavities as mentioned by these authors are also lacking; they seem to be an artefact caused by re‐expanding dried fruits. The seed coat is not sclerified. The fruitlets of Amborella differ from externally similar fruits or fruitlets in other basal angiosperms, such as Austrobaileyales or Laurales, in their histology. © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 148 , 265–274.     

Embryology and fruit development in four species of Pistacia L. (Anacardiaceae)

Pistacia atlantica, P. palaestina, P. lentiscus and P. saportae , were found to have great similarity in their embryology and fruit development. The anatropous, pendulous and crassinucellate ovule was initially unitegmic; later, the integument split close to the micropyle, forming a partial second integument. After anthesis there was a development of a hypostase and an obturator. The development of the Polygonum-type embryo sac followed division of a megaspore mother cell, giving a tetrad or triad of megaspores. The functional megaspore was the chalazal one. The ovary developed into a mature pericarp after anthesis, even when pollination was prevented, and before the zygote divided. Therefore, the fruit can be parthenocarpic. The ovule started to grow after initiation of embryo development until it filled the cavity within the pericarp. The zygotes were dormant for 4–18 weeks after pollination. In P. saportae reproduction became arrested during the development of the embryo sac; only very few abnormal embryos were found. No fixed pattern of embryo development could be discerned. The endosperm was initially nuclear, becoming cellular when the embryo started to develop. The seed coat was derived from the integument and the remnants of the nucellus.     

Extensibility of pericarp tissue in growing citrus fruits

The tensile force existing in the pericarp of a growing citrus (Citrus sinensis) fruit 17 to 19 centimeters in circumference was sufficiently high to cause a 3% shrinkage of the pericarp when it was excised. When a fruit was cut along the equator to the central axis, shrinkage of the pericarp resulted in the formation of a wedge-shaped gap at the cut. Stretch modulus of the pericarp was determined by measuring the force required to stretch excised strips of tissue to 1% longer than their excised length. Measurements were made on successive layers of pericarp tissue 5 millimeters wide and 1 millimeter thick taken from the fruit equator. All layers required more force for extension at lower temperatures and high water potentials than at high temperatures and low water potentials. The stretch modulus ranged from 0.88 to 2.16 kilograms per square millimeter depending upon the layer, temperature, and water potential. The inner layers, consisting primarily of mesocarp, had stretch moduli only 60 to 70% as great as the outer layer which consisted of exocarp tissue. Measurements of the stretch modulus of tissues from the pericarp support the hypothesis that changes in the tension existing in the pericarp depend upon conditions in the pericarp and are not related to changes in volume or pressure in the juice vesicles.     

杧果果实的扫描电镜观察

果成熟果实呈黄色长卵形是典型的核果。其外果皮呈革质,中果皮呈肉质,内果皮呈骨质表面有纤维。内果皮包被着种子形成一个大而扁的核。种子扁形,种皮很薄,子叶呈多胚或单胚。通过对果果实结构各个层次的扫描电镜观察,为其贮藏保鲜及繁育栽培工作提供一些科学依据。