On Some Regularities of One-Seeded Fruits Evolution
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Journal of Life Sciences 9 (2015) 511-520 doi: 10.17265/1934-7391/2015.11.001 D DAVID PUBLISHING On Some Regularities of One-seeded Fruits Evolution Тatyana I. Kravtsova Komarov Botanical Institute RAS, Saint Petersburg 197376, Russia Abstract: Anatomical structure of differently originated seed envelopes in one-seeded indehiscent fruits of Urticaceae and Asteraceae members is studied using light and scanning electron microscopes. It was found that in anthocarps and involucrate fruits of both families the relations between the primary (pericarp) and secondary fruit envelopes (perianth and/or involucre) were composed under complexification (union) type, and not as substitution. Numerous examples of non-homologous resemblance in fruit envelope structure indicate a high degree of adaptability of certain histological types, recurring on a different morphological basis in different phyletic lines within a family. These tissue complexes represent widely occurring types of the pericarp (Utricaceae) or pericarp and seed coat tissue union (Asteraceae). This evolutionary repetition or pseudocyclic resemblance is apparently another common regularity of one-seeded indehiscent fruits evolution in addition to those enumerated in general by Zohary (1950). Key words: Asteraceae, Corymbium, Urticaceae, accessory envelope, fruit, non-homologous resemblance. 1. Introduction highly organized than other fruit types, because functional correlations between their elements are Molecular-phylogenetic investigation of stronger [11]. As was noted by Zohary [9], it is Angiospermae [1], detailed studies of gynoecium especially intriguing to trace further evolutionary structure [2-4], paleobotanical findings [5], changes of fruits in those groups where monospermy examination of morphological and anatomical fruit has become a fixed characteristic. He and the characters in many extant taxa have broadened our followers (e.g., Claßen-Bockhoff [16]) found, that knowledge about fruit structure summarized earlier by similar morphological transformations occurred Roth [6], especially in basal angiosperms. The origin subsequently: aggregation of fruits into dense of a carpel and the very first steps of fruit evolution infructescences (aggregates) corresponding to remain, however, uncertain questions [2, 4, 5, 7]. The functional units, differentiation of fruits to the notion that seemingly stayed changeless is that infructescence (heterocarpy), their numerical one-seeded indehiscent fruits (pericarp fruits [8]) of reduction up to one, aggregation of mono- or living taxa generated by the reduction of apocarpous oligomerous dense infructescences into secondary and coenocarpous fruits, what often followed by aggregates. Involvement (incorporation) of integration with flower and inflorescence. According extracarpellary flower organs in fruit formation is to the concept, this fruit represents an ultimate stage of common trend as well [6, 8, 10, 17-19]. one of general trends of fruit evolution [6, 8-12]. Deep Our observations have shown that those families are morphological differences of fruits, their protective of particular interest for study fruit evolutionary trends, and dissemination functional dissimilarities are related in which one can trace the transition of pericarp fruit to poly-/monospermy and dehiscence/indehiscence (in the sense of Pijl [8]) to fruit-anthocarp (in sense of [10, 11, 13-15]. One-seeded indehiscent fruits are Spjut [20]); alias, where fruit as the system considered sometimes as very integrate systems, more “fruit-seed” [18] transforms into the system “infructescence-(flower)-fruit-seed”. In newly Corresponding author: Тatyana I. Kravtsova, Sc.D., leading research scientist, research fields: botany, evolutionary arranged anthocarps or involucrate fruits the morphology, comparative anatomy of higher plants. E-mail: redistribution of protective and dissemination [email protected]. 512 On Some Regularities of One-seeded Fruits Evolution functions starts again between different seed using scanning electron microscope Jeol JSM-35C envelopes, and partial transference of functions to and light microscope AxioImager А1 (Carl Zeiss) secondary envelope occurs. equipped with digital imaging AxioCam MRc5 and The goal of this work was to study specific software Zen 2011. character of evolutionary transformation of this fruit 3. Results and Analysis type by the example of two families—Urticaceae and Asteraceae (tribe Corymbieae), the question on 3.1 Urticaceae structure and contribution of differently originated Fruits in the nettle family develop from a seed envelopes to fruit formation was in the focus of pseudomonomerous gynoecium [22, 23], the ovary is attention. unilocular with single orthotropous ovule, attached 2. Materials and Methods basally or subbasally. In the Urticaceae several Fruit structure in 253 species and some evolutionary trends may be distinguished in fruit infra-specific taxa from 44 genera of the Urticaceae development [24, 25], among them partial and in only few Asteraceae, three Corymbium species, transference of pericarp functions to the accessory was examined using light and scanning electron fruit envelope—fruiting perianth and involucre, the microscopes. The study was based mostly on fruits development of winged and fleshy drupe-like from herbarium specimens from LE, K, L, MO, VLA, disseminules, dense infructescences, representing PE, and some others, partly on material from botanical functional units and bearing resemblance with true gardens (seed exchange), carpological collection of fruits. The last is often accompanied by the Komarov Botanical Institute (St. Peterburg, Russia), development of more or less fleshy or juicy fruiting greenhouses of Komarov Botanical Institute, and perianth, promoting fruit aggregation, fleshy Main Botanic Garden of Russian Academy of receptacle (Pipturus, Nothocnide, Procris) or Sciences (Moscow, Russia), and were collected in inflorescence axes (Gyrotaenia). It should be noted nature—St. Petersburg vicinity, the North Caucasus. that just obtaining of this qualitatively new accessory For anatomical study, fruits were first placed in a fruit envelope that resulted in emergence of the new mixture of water, 96% ethanol and glycerol in equal diaspore with higher organization level; it increased proportions. Material was fixed in 70% alcohol, and fruit evolutionary plasticity, supplied the diversity of for thin sections in 3% glutaraldehyde and 2% dissemination modes and fruit polymorphism Osmium tetroxide. Cross sections 12 and 24 µm thick (heteroanthocarpy, genetic polymorphism) in the were made in the middle part of fruits using freezing group. Together with transformed perianth and the microtome, and histochemical studies were carried out involucre, arising as the result of oligomerization of to determine lignin (with phloroglucinol and sulfuric inflorescence in representatives of Forsskaoleae, acid) and cutin (with Sudan IV). Cross and Parietarieae and Boehmerieae the fruits sometimes longitudinal paraffin sections (12 µm thick, stained acquire lignified, more or less thick cover (Hemistylus with gentian-violet and orange or Safranin and alcian boehmerioides, Neodistemon indicum Parietaria blue) were made on rotary microtome following judaica, P. cretica, certain Pouzolzia species). standard procedures [21], and semi-thin sections (2-3 It was found in the result of comparative anatomical µm thick, stained with toluidine blue) of material examination [25-27] that few layered, in many genera embedded in Epon-Araldite epoxy resin were prepared of the same type, highly specialized pericarp, often on Reichert Ultracut R ultramicrotome. Observations strongly mineralized, has been formed in parallel with were carried out and photomicrographs were taken foregoing processes of fruit complication and On Some Regularities of One-seeded Fruits Evolution 513 aggregation. Strongly reduced seed coat in Urticaceae genera simultaneously with monotypic structure of the is represented by tanniniferous membrane, brown or pericarp. Firstly attached to fruit, membranous and greenish due to chloroplasts, sometimes (Urtica) easily removable (certain Boehmeria species), it adhering to the pericarp. Two-three strongly subsequently becomes inseparable and structured, compressed layers may be distinguished in the seed acquires histological differentiation. Fleshy or juicy coat, the exotesta is often perforated, while the fruiting perianth may develop in parallel in different endotegmen tanniniferous. The presence of tannins genera irrespective of pericarp structure; dry and endotesta cells provided with lignified thickened structured fruiting perianth is connected commonly anticlinal walls (Boehmerieae) indicates to with hard mineralized pericarp. maintenance by the seed coat of protective role to It is found that this evolutionary fruit some extent. Another function may be assumed for it, transformation in the Urticaceae is not usually connected with respiratory metabolism during early accompanied by displacement of its primary seed development: testal aerenchima is formed in envelopes (Figs. 1A-1G). It is true only in respect of outer integument, and holes (intercellular spaces) involucre, that may take place of thin fruiting perianth appear in the exotesta at the stage of globular embryo. to the large extent, and of thin and considerably The degree of seed coat development is not in reverse reduced pericarp in the case of lignified or vigorous correlation with pericarp structure, as it was found, involucre (Parietaria