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Genus Salsola of the Central Asian Flora; Its Structure and Adaptive Evolutionary Trends (中央アジアの植物相salsola属の構造と適応進化の傾向）

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Genus Salsola of the Central Asian Flora; Its Structure and Adaptive Evolutionary Trends (中央アジアの植物相salsola属の構造と適応進化の傾向） Genus Salsola of the Central Asian Flora; Its structure and adaptive evolutionary trends (中央アジアの植物相Salsola属の構造と適応進化の傾向） 2008.9 Tokyo University of Agriculture and Technology Kristina Toderich （クリスティーナ・トデリッチ） 1 Genus Salsola of the Central Asian Flora; Its structure and adaptive evolutionary trends (中央アジアの植物相Salsola属の構造と適応進化の傾向） 2008.9 Tokyo University of Agriculture and Technology Kristina Toderich （クリスティーナ・トデリッチ） 2 Genus Salsola of Central Asian Flora: structure, function, adaptive evolutionary trends, and effects to insects Table of Context CHAPTER 1. Introduction 1.1. Introduction………………………………………………………………………. 1 1.2. Order Centrospermae and taxonomic position of genus Salsola………………… 4 CHAPTER 2. Material and Methods 2.1. Desert ecosystems of Uzbekistan ……………………………..……………….. 13 2.2. Ecological and botanical characteristics of Asiatic genus Salsola………………. 28 2.2.1. Botanical and economic significance of Central Asian Salsola representatives… 28 2.2.2. Plant material (examined in the present study)…………………………………… 31 2.3. Methods of investigation…………………………………………………………. 35 CHAPTER 3. Floral micromorphology peculiarities of micro-and macrosporogenesis and micro-and macrogametophytogenesis 3.1.1. Biology of and sexual polymorphism of flower organs at the species and populational levels……………………………………………………………… 50 3.2.Anther wall development, microsporogametophytogenesis and pollen grain formation…………………………………………………………………… 58 3.3.Morphology, ultrastructure, interspecies variability and quantitative indexes of pollen grain……………………………………………………………………….. 64 3.4.Ovule morphology, peculiarities of megasporogametophytogenesis and development of embryo sac……………………………………………………… 89 3.5. Floral morphology and taxonomic relations among species of genus Salsola…... 94 CHAPTER 4. Processes of flowering, ecology of pollination, fertilization, embryo development and their adaptive significance…………………………………102 CHAPTER 5. Morphology and anatomy of fruits 5.1. Species diversity and polymorphism of carpological characters…………………... 111 5.2. Seed storage, type of dormancy and ecology of germination……………………… 126 CHAPTER 6. Vegetative floral organs micromorphology, ion/translocation mechanisms 3 and photosynthetic pathways 6.1. Diversity of morphology of trichomes, hairs and perianth stomata in some taxa of Salsola………………………………………………….………………………….129 6.2. Salt accumulation, silicification, wax deposit in the associated epidermal structures of the inflorescence bracts and bracteoles……………..…………………………. 136 6.3. The occurrence of C3/C4 photosynthesis in the leaf-like organs of Salsolas under Kyzylkum Deserts conditions……………………..……………………………. 152 6.4. Sexual reproduction, photosynthetic pathways and biochemical linkage…….... 160 CHAPTER 7. Isoenzymatic and karyological studies of some Salsolas species... 163 CHAPTER 8. Adaptive strategy and evolutionary trends in Asiatic genus Salsola its taxonomic relations within Centrospermae -Reproductive strategies of Haloxeric Asiatic Salsola species and its adaptive significance-…………….. 172 General Conclusions.………………………………………………….……………… 177 ACKNOWLEGEMENT.…………………………………………….……………… 183 REFERENCES.………………………………………………….…………………… 184 4 CHAPTER 1. INTRODUCTION 1.1 Introduction Uzbekistan, in the Central Asian region, is a landlocked country covering 447,400 square kilometers. Two-thirds of its land consists of steppe, desert, and semi-desert areas with the remainder as fertile valleys with abundant natural resources: oil, natural gas, gold, polymetals and so on. Dramatic political changes after the collapse of the Soviet Union have impacted all areas of human life, including the relationship between humans and natural resources. Over-exploitation of natural resources particularly through unsustainable agricultural practices is central to the issue. Some of the adverse actions of considerable importance are irrigation, alteration of natural rivers, mining, drilling, oil and mineral exploration, poor infrastructure development (roads, markets and railways), urban development, fuelwood collection, over-stocking and overgrazing by livestock. The poor management or destruction of appropriate native woody-shrub/ dwarf shrub vegetation in many arid areas of Uzbekistan is leading to land degradation with the development of wind and water erosion, preventing natural regeneration of rangeland plants. Shifting sands are one of the major problems of land degradation in the Kyzylkum Desert. Shifting sand dunes are almost devoid of vegetation and make up about 0, 5 million hectare (Kharin, 1995). Grazing and trampling by livestock, agricultural use, and mining industries destroy the vegetation cover and enhance sand mobility. Sand dune movement is a threat to irrigated farmlands, villages, railways, highways and others infrastructures. The leading factors of anthropogenic degradation of cover vegetation in the Kyzylkum deserts are: a) agricultural land use (grazing, cutting and up-rooting of shrubs and undershrub), b) hay making and c) industries impact (mining, construction activity, transport, etc). Salinization is one of the major ecological and production problems currently facing the agricultural sector in arid and semiarid areas of the Central Asian countries. Recent use by the former USSR of the major rivers of Central Asia (Amudarya, Zerafshan, Syrdarya) for the production of exports, including cotton, oil, and minerals, has resulted in rising water tables, waterlogging, and the well known ecological disaster of the saline lands around the receding Aral Sea. In particular, secondary salinization (human caused) is increasing rapidly and crop production under these conditions is becoming less sustainable in this region. Drought and salinity are capable of having a far greater effect on food security in Central Asian than in other areas. Besides salinization, contamination by heavy metals and chemical compounds released by agriculture, uranium, oil and gas industries has been frequently reported for Kyzylkum sandy Desert (Walter and Box, 1983; Goldshtein 1997; Solodov 1998; Toderich et al 2001, 2002, 2004a, b; Tsukatani et al 2001). Initial explorations of the natural plant cellular mechanisms affecting the 5 bioremediation of elemental and/or organic pollutants suggest great promise for the use of desert plants in large-scale environmental clean-up efforts. Factors that relate to take-up of chemical compounds, such as tannins, nitrates, metals, and oxalates (some of which may be toxic for plant development) have not been adequately studied for the arid flora of Central Asia. Native desert plants, however, have the advantage of being highly adapted to the hyper-arid climatic and edaphic contaminated conditions. There is firm evidence that conditions within the core areas of Kyzylkum Desert are getting worse and that some remedial effort is warranted to protect the biodiversity, urban settlement, resource extraction and communication links. In this context the native Salsola (from order Centrospermae, Chenopodiaceae family) species have the advantage of being highly adapted to the local deserts edaphic and climatic conditions, which are characterized by having a strong seasonal factor. Salsola plants, which are an innovative pioneer species for range regeneration of the sandy Central Asian deserts, serve as an excellent model to study sexual reproduction, seed dormancy and native seed bank establishment for arid fodder plants. Knowledge about Salsola ”plant functional reproductive system” can be useful in the evaluation and prediction of sexual reproduction, seed formation and ecology of seed germination techniques, especially for the conservation and restoration of natural saline/sandy desert ecosystems in Uzbekistan. The conservation and protection of gene pools of native arid and semiarid Asiatic flora is basic for understanding the influence of environmental factors on morphoembryological processes, as well as revealing of their plasticity and tolerance under stress conditions. Plant reproductive strategies, based on morphological, structural, physiological and biochemical features for different ecological groups of chenopods of the desert Central Asiatic flora, are not well documented in the literature (Butnik 1981; Toderich et al., 1995, 1998, 2002, 2004a; P’yankov et al., 1997; Wolfe, 1998). Traditionally descriptive anatomical and embryological approaches were based mainly at the families or generic levels. However several key features are unreliable since polymorphism within taxa often exceeds interspecific boundaries. In the present work our proposed concept of “plant functional reproduction system” (PFRS) is based on common plant traits both structural and functional, that are responsible for similar responses to environmental conditions and that have dominant effects on major ecosystem processes. Groups of plant species with common structural features can form a “functional plant group”. Plant traits used for the identification and level of adaptation of species include important structural embryological parameters of floral organs, pollen biology and morphology, effective pollen control and pollination system, structural fruits traits, seed set and seed germination characteristics connected with reproductive plant functional activities (Wallace et al, 1968; Werker 6 and Many, 1974; Young et al., 1983; Yamaguchi, 1990; Takeno et al. 1995). The fertility benefit of reproductive assurance for arid plants has not been examined in the light of their seed production. The reproductive plant functional hypothesis resolves this disparity
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