More Reveals Of Xerophytic Species

Fig. 12.1 Distribution characteristics of trace elements in various desert taxa

Halopeplis pygmea, Amaranthus retroflexus, Limonium sogdianum, Sonchus maritima Puccinella scleroides, Sorghum bicolor, Peganum harmala, Haloxylon aphyllum, as well as annual and perennial species of the genus Salsola. These pioneer plant species were growing well on mined areas despite unfavourable conditions such as extreme pH, high salinity and phytotoxic levels of several elements.

4 Salt Accumulation, Silicification, and Wax Deposition Associated With Epidermal Structures of Flower

Desert plants successfully growing on metalliferous or salinized soils tend to accumulate the highest ion concentrations in epidermal and subepidermal tissues, including various glandular structures of bracts/bracteoles and perianth segments. Salt glands with varying degree of specialization are actively involved in the elimination of solutes and mineral elements from the surface of the vegetative organs. These are very common in the desert plants of Central Asia. Excretion occurs predominantly on the adaxial surface and is uniformly localized along the lateral walls of the grooves (Figs. 12.2). Salt glands morphology vary in different genera. These can be sunken, semi-sunken or located above the epidermis as in the majority of chenopods and gramineae species. In the latter glandular structures are usually bicellular, comprising a basal and cap cell. Slight variations in morphology of the basal and cap cells of glandular hairs have been observed mostly in the annual and other species such as that of genus Salsola, Aeluropus litoralis, Tamarix hispida, and Eremopyrum orientale, which appear to be related to their efficiency of salt secretion (Toderich et al. 2003, 2008).

Our findings showed that epidermal vesicles and papillae in desert species (Figs. 12.2 and 12.3) have a large bladder cell attached to a stalk composed of one or more cells that in turn is attached to an epidermal cell. Comparative study of two annual taxonomically close related Salsola species from steppe soils of Europe (Poland) and Kyzylkum metalliflerous/salinized sands revealed that salt secretion become prominent and salt glandular structures are formed abundantly only when plants are exposed to high contaminated environments. Under such conditions an evident increase in succulent bracts is a consistency met within Kyzylkum chenopods.

The vesiculate hairs of some annual Salsola from Central Asia are considerably involved in the cellular salt secretion. According to Luttge (1971) this might not be taken strictly as a secretor process, because these trichomes are considered

Fig. 12.2 The morphology of Vesicular-and short peltate trichomes on bracts of Salsola pestifer (Buchara ecotype)
Fig. 12.3 Epidermal surface view of mature bracts of Salsola pestifer (Buchara ecotype). Glandular structures have a strong localization, especially on adaxial side, which is mostly exposed to environmental impact. X 750

as salt glands and their function is obviously a specialized mechanism for the removal of salt from the leaves. The emission of salt from these vesiculated hairs is apparently the result of the rupturing and collapse of bladder cells (Gamal 2005; Ottenhof et al. 2007). The presence of papillae on the epidermal cells of S. praecox, S. iberica and S. pestifer, with thick outer walls, cuticle and submerged stomata seemingly protect assimilatory organs against excessive transpiration. C4- herbaceous annual Salsola species differ in the morphology (head shape-mainly clavate or capitate or also in the number of constituent cells composing their stalk) of salt-glands/trichomes and their density on the epidermal surface. Variation in the density of salt glands/trichomes is believed to be mainly due to the effect of stress under desert environmental conditions and even pressure from herbivory (Wahid 2003). These parameters potentially could be used as distinguishing characters between different ecological halophyte groups. For instance the Climacoptera complex has unicellular non-glandular trichomes or hairs, smooth or micropapillate (warted surface), whereas the surface of bracts/bracteoles of many dry/sclefiried Salsola species have an undulating epidermal surface with numerous salt glandular structures and tall adaxial ridges alternating with deep grooves. On the ridges of annual Salsola species we found various papillae and prickle hairs, as well as secreted salts, which appear as crystals. Crystalline deposits were more abundant on the adaxial surface because of higher gland frequency (Fig. 12.4a, b).

It has also been noted that occurrence of calcium oxalate crystals was almost absent in root and stems. An abundance of these crystals was described in the tissue of the seed coats of many xero- and euhalophytes.

Salt glands usually are globose or club-shaped and readily distinguishable from unicellular papillae and sharp-pointed prickles, an ornamented, porous cuticle overlies the epidermis, cuticle is distinctly thicker over the area that adjoins basal and epidermal cells than over the cap or other parts of the epidermis. The cuticle is separated from the outer cap cell wall, resulting in the formation of a salt collecting Chamber (Fig. 12.4b) or cuticular cavity which is similar in the species of Salsola (both annual and perennial), Aeluropus litoralis, Eremopyrum orientale, Spartina,

Fig. 12.4 a The adaxial surface of bracts of S. iberica with ridges and salts which appear as crystals. b SEM microgrphs showing the patterns of crystaloid structure in the bract tissues of S. orientalis

Cynodon and Distichlis (Thomson 1975) and probably represents a temporary collecting compartment where secreted salts accumulate prior to elimination from the leaf. The ions seem to be compartmentalized in small vacuoles and transported to the cuticular cavity, prior to exclusion from the vegetative and reproductive organs either through cuticular pores or by rupture of the cuticle (Yordoan and Kruger 1998; Naidoo and Naidoo 1998; Rozema and Riphagen 2007).

An unusual type of salt glandular structure has been described for Salsola car-inata where the terminal cell(s) always ends bluntly. On top of the stalk cell, extremely thin-walled cells form a single originally ornamented ring, while the thick cuticle of the stalk cell remains as a cylindrical scar (Figs. 12.5 and 12.6a, b).

Cross-sections of bracts and bracteoles of many Salsola species show that different tissues like swollen epidermal cells (in all species), large-celled hypodermis and water bearing parenchyma carry out water and salt-accumulating functions. Size, shape and/or their density should be recognized by the location and deposits of

Fig. 12.5 a Salt gland of Salsola paulsenii comprising flask-shaped basal cell, dome-shaped cap cell and raised cuticular chamber. b Micromorphology of glandular hairs of Salsola sclerantha and wax-epicuticular inclusions partially surrounding it
Fig. 12.6 a SEM micrograph showing surface features and morphology of non-glandular, unicellular hair of bracts in Cimacoptera lanata. b Untypical morphology of salt land, occurring on the epidermal bract's surface of Salsola leptoclada (Central Kyzylkum ecotype)

Fig. 12.7 a Cross-section of succulent bract of Salsola praecox; central part is occupied by 3-4 layers of water-storing parenchyma cells with small salt crystals 10 x 60 (1 ^k). b Anatomy of bract tissue of Salsola arbusculiformis. Different types of crystals in the subepidermal salt-storage cells 10 x 60 (3.0 ^k). c Cross section of anther in Salsola arbuscula. The salt ions location in pollen grains (male gametophyte) is absent 10 x 60 (1 ^k). d The fluorescent microscopy image of bract of S. arbusculiformis with the location of salt/ions in it 10 x 60 (1 ^k)

Fig. 12.7 a Cross-section of succulent bract of Salsola praecox; central part is occupied by 3-4 layers of water-storing parenchyma cells with small salt crystals 10 x 60 (1 ^k). b Anatomy of bract tissue of Salsola arbusculiformis. Different types of crystals in the subepidermal salt-storage cells 10 x 60 (3.0 ^k). c Cross section of anther in Salsola arbuscula. The salt ions location in pollen grains (male gametophyte) is absent 10 x 60 (1 ^k). d The fluorescent microscopy image of bract of S. arbusculiformis with the location of salt/ions in it 10 x 60 (1 ^k)

salt/ions in specific (salt- storage) cells. The fluorescent microscopy studies on the displacement of salt ions from the floral organs of some Salsola species reveals an abundance of mineral ions in the tissues of sterile organs of flower like sepals and anther connective cells. However ion dislocation has never been observed in male-and female gametophytes or in the embryo tissues (Fig. 12.7a-d).

Occurrence of calcium oxalate crystals in the leaves and seed coats of some plants has been described by Fuller and McClintock (1986). It has been suggested that concentration of oxalate crystals is almost absent in the root and stems. The presence of crystals in the outer covering of seeds may play a role in changing soil pH, thereby providing a more favourable condition for plant survival.

Structurally, SEM studies revealed a high diversity in the micromorphology of epicuticular wax (epicuticular secretion), mostly occurring as specific crystalloids (epicuticular wax crystalloids) on the plant surface of desert plants as proposed by Barthlott et al. (1998). Cuticular wax partially covers the mature prickle-hairs, papillas and long cells of outer epidermis of bracts/bracteoles of some perennial Salsola species as is shown at Fig. 12.8a,b.

Their nature and molecular organization of such wax deposits is still unknown for desert plants. The chemical composition of these waxes has been given at length by Barthlott (1994) and Barthlott et al. (1998). However, there are still contradictory opinions concerning waxes deposition. Earlier workers suggest that waxes could be exuded to the outer cuticular surface through pores, while Mahllberg (1991) suggests that there is excretion through lamellate regions onto the cuticle. Glandular trichomes in such case enhance capacity of plant to accumulate large quantities of

Fig. 12.8 a Scanning micrograph of Salsola orientalis bract epidermal surface with various salt crystalloids (or epicuticular inclusions) on it. b Silicon X-ray distribution image of mature inflorescence bracts in Eremopyrum orientale (Poaceae) X 3000

volatile components and transport these to the cuticular surface for vaporization from the gland surface.

A comparative developmental study of floral organs of various chenopods and graminceous species revealed that Si accumulation was greatest on the adaxial tri-chomes of inflorescence of Eremopyrum orientale, Bromus tectorum and Aeluropus litoralis, collected from highest contaminated areas of the Bukhara oasis. The localization of small siliceous particles on the inflorescence bracts of Eremopyrum orientale is concentrated mostly on the surface of epidermis around stomata. Crystalloid types in Salsola taxa are characterized by uniformly distributed small irregular-shaped platelets which occasionally have a parallel orientation around the stomata. In some chenopod species platelets occur in clusters too. A similar sili-cification process associated with trichomes and other epidermal structures of the inflorescence bracts was described for Phalaris canariensis. It is said that the silifi-cation may be synchronized with the deposition of wall substances, such as lignin, suberin and phenols (Sangster and Wynn Parry 1981). Silicon deposition patterns and localization in bracts has been described for different groups of flowering plants (Sangster et al. 1983; Hodson et al. 1983; Rufus et al. 2007).

Electron microscopic X-ray analysis of secretion products from the salt glands in different representatives of Salsola shows a localization of variety of mineral elements and ions. Prismatic crystals secreted by glands primarily contain cations Na, K, Ca, and anions Cl, SO4, carbonate, although other ions such as Mg, Si, Sr were also detected. These findings require further studies on a wider range of plant materials with respect to structural and genetic variation and their relation to bioremediation of contaminated desert ecosystems.

We can conclude that sandy and saline soils contaminated with Cd, Sr, Cu, Fe, Ni, Mn, Cr, Pb, Zn, and various toxic salts and organic pollutants are colonized by plant species that develop strategies for avoidance and/or tolerance to metals. One possible avoidance strategy is preventing uptake of potentially toxic metals, especially into the reproductive organs like pollen grains and embryo. Although tolerant plants seem to restrict salts and metal uptake to varying degrees this mechanism still has not been strongly analyzed in arid vascular plants. It was found that salt (minerals and ions) accumulating glands are mostly common in families Poaceae, Tamaricaceae, Chenopodiaceae, and Frankenaciaceae, and occurr only in a few scattered species in the families Plumbaginaceae, Zygophyllaceae, Fabaceae, and Lamiaceae. Many species of these families are known to have glandular structures, but further investigations are needed to determine their secretion products.

5 Diversity in Trichomes, Hairs and Salt Glands (SEM)

Trichomes are highly variable appendages of the epidermis including glandular (or secretory) and nonglandular hairs, scales, papillae etc., varying widely in structure within larger and smaller groups of plants and are sometimes remarkably uniform, and may be used for taxonomic purposes. The glandular forms are structures on the plant leaf/perianth surface, usually in direct contact with surroundings, playing a defensive role against herbivores and pathogens, in the salt secretion, plant pollination and other interactions between plants and environments; due to their morphology and production of different chemical products. Still there is neither a satisfactory nor well-accepted classification of trichomes for higher plants (Behnke 1984). The importance of the micromorphology and distribution of glandular trichomes for the taxonomy of some species and subspecies requires a reconsideration, because morphology and ultrastructure can be used as a valuable marker for the evolutionary level of the taxa. The pronounced variability of glandular structures can also be related to phenotypic responses to salinity or contaminated environments. These have been used in the delimitation of the sub-families of Chenopodiaceae and the categories are fairly homogenous with regard to trichome type. Carolin (1983) has studied the trichome morphology and its classification within Chenopodiaceae and Amaranthaceae. The morphological traits of trichomes and/or hairs provide a key for easier identification and delimitation of the closely related taxa in different flowering plant groups. The herbaceous Central Asian halophytes; well known in the pasture economy of Uzbekistan as "solyanki"; differ from European taxa in the morphology of salt-glands/trichomes (shape of their head, mainly clavate or capitate and its density). An abundant papillae, prickle hairs and salt secretion between ridges on the surfaces of bracts/bracteoles of annual Central Asian Salsola species reveals that frequently salt glands are globose or club-shaped and readily distinguishable from unicellular papillae and sharp-pointed prickles. These parameters can be used as discriminating characters between different ecological variants of Salsola group. Variation in the indumentum density is believed to be mainly due to the effect of stress under desert environmental factors and/or even herbivory pressure.

An assessment of the validity of trichome characters and their morphological diversity under harsh desert and contaminated environments was evaluated. The main trichome types for the Central Asian species of Salsola are schematically shown in Fig. 12.9.

S. orientalis


Fig. 12.9 Diversity of trichome morphology (SEM) in some species of genus Salsola: S. orientalis and S. incanescens from section Caroxylon are clearly separated from all other species of genus Salsola by the development of branched and dendric trichomes; malpigian type of hairs are characteristic of S. gemascens

S. orientalis

S. aperta


Fig. 12.9 Diversity of trichome morphology (SEM) in some species of genus Salsola: S. orientalis and S. incanescens from section Caroxylon are clearly separated from all other species of genus Salsola by the development of branched and dendric trichomes; malpigian type of hairs are characteristic of S. gemascens

The species of Central Asian genus Salsola exhibit two unicellular trichome types as described earlier by Carolin (1983) in the families Chenopodiaceae and Amaranthaceae. Using the indumentum characters we found that the Salsola species examined by us can be allocated to different sectional groups. A nonparametric analysis of variance of the densities of unicellular/multicellular trichomes on the surfaces of bract/bracteoles, as well as number of cells composing the stalk of mul-ticellular trichomes revealed that the trichome characters studied possess different values for each species and these might be valuable when identification is impossible using macromorphological parameters. Trichomes have highly variable appendages of the epidermis including glandular or secretory and nonglandular hairs, scales, papillae. Trichomes of Central Asian Salsola species have been classified by us into a few morphological categories such as; hairs, which maybe unicellular or multi-cellular; glandular or nonglandular; scales or peltate hairs; water vesicules, which represent enlarged epidermal cells. Glandular hair-a trichome having a unicellular or multicellular head composed of secretor cells, which is usually borne on a stalk of non-glandular cell varying in the degree of differentiation.

For majority of species of Salsola-non glandular clothing trichomes, unbranched, uniseriate, multicellular are composed of one or two basal epidermal cells and one or six cells are arranged in one row. Their surface is usually covered by cuticular micropapillae lacking basal part of the trichomes. The glandular structures are usually bicellular, comprising a basal and cap cell, and are referred to as salt glands, trichomes or microhairs.

Based on the analysis given above we propose the following classification of glandular structure for genus Salsola:

a. Papillae, the most simple and common type of glandular hairs in the genus Salsola scales, huge or sessile glands that can be found in species of section Salsola and sect. Arbuscula consists of a short stalk of two parallel cells and multicellular glands, often cuticula is removed;

b. Peltate trichomes with one basal cell, one stalk cell, and glandular head; the subcuticular space is remarkably large;

c. Unbranched, short glandular hairs, stalk bi-or multiceseriate, gland spherical, basal biseriate with two very short cells and a few secretory cells;

d. Long capitate trichomes, which have usually one (sometimes two) basal cells; the stalk composed of one to four cells (the upper one is often shorter and marked as neck cell and one cell head; sometimes with small subcuticular space);

e. Simple two-armed, unbranched glandular hairs, stalk cells are usually thin-walled, these types of Glandular hairs could only be found on the bracts/bracteoles and tepals of S. gemmascens (sect. Malpigila);

f. an unusual type of salt glandular structure was described for S. carinata, the terminal cell(s) for many Salsola species always end bluntly on top of the stalk cell, extremely thin-walled cells form a single originally ornamented ring, while the thick cuticle of the stalk cell remains as a cylindrical scar.

The unicelled and stiff trichome on multicellular base is one of most frequently found type within genus Salsola. This type of uniseriate smooth trichomes are mostly common found in the species of section Salsola and Physurus and have no noticeable differences in texture between the body and base, which is more or less bulbous. However, it remains to be explained if long (as in the case of species of section Physurus) and short (described for S. paulsenii, S. praecox, S. pestifer, S. iberica) trichomes represent two different kinds or two different developmental stages of the same trichome. Dense epidermal-cell protrusions or few-celled of well developed smooth trichomes, which were described for some species of sect. Physurus, obviously, indicate that these species are tolerant to extreme dry and saline habitats.

Our results showed that Central Asian annual species, especially from sect. Salsola subsec. Kali can be clearly separated from the annual species of the same section from Europe, not only on the basis of morphology, but also by the density of unicellular trichomes on both bract/bracteoles surfaces. Micropapillate unicellular trichomes are highly specific to S. paulsenii, S. praecox, S. pestifer, S. iberica. The closely related annual European species of section Salsola subsec. Kali in particular S. ruthenica and S. kali are similar with Asian annual Salsola species, except for the density of glandular trichomes on the bract/bracteoles surfaces. S. ruthenica and S. kali possess smooth bract/bracteoles surface or with a presence of slightly developed papillae-a soft protuberance structures. This probably indicates a co-species relationship between the Asian and European species of genus Salsola.

Although an abundant development of various types of trichomes within desert Asian Salsola species might be well correlated with the desert ecological factors. Wide morphological variations are exhibited by the species of sect. Cardiandra and Belanthera, which mostly possess both uni-and multicellular trichome types (bladder cells-structural organization) which are usually globose or club-shaped and readily distinguishable from unicellular papillae and sharp-pointed prickles.The 2-armed or detached smooth trichomes called 'Malpigilian hairs' seem highly specific to species of sect. Malpigila, while vesicular and various glandular structures are best represented in the species of sections Cardiandra and Belanthera.

It has been observed that in some cases an accumulation of high concentration in the vacuole of terminal cells of bladder trichomes are released probably by rupture of the cytoplasm and cell-walls (Thomson et al. 1988). In such cases the collapsed cell gives the characteristic mealy appearance of the epidermis in many Chenopodiaceae. Therefore with the help of morphological characters; mainly related to epidermal structures (by SEM analysis); we find that the Salsola species complex could in fact be divided into two groups: species with salt-producing tri-chomes/hairs and salt-accumulating (with specific salt/storage cells) plants. This stresses the fact that different mechanisms and strategies for the sequestration and regulation of the salt ion concentration in the plant tissues are operated in the stem and leaf succulent halophytes and in the recreto-and pseudohalophytes of the Kyzylkum flora. The ability of some desert chenopods to accumulate significant amounts of nitrates and/or oxalates has been reported b y several investigators notable among them being (McWorter et al. 1995; Sandquist and Ehleringer 1997; Judd and Ferguson 1999; Butnik 2001a, b; Wojnicka-Poltorak et al. 2002).

The natural plant-cellular mechanism of salt/metal removal and tolerance presented here shows that more detailed studies are needed for a development and testing of more valid hypothesis regarding the adaptations required for colonization and survival of plants, growing under extremely harsh and simultaneously contaminated desert environments.

It is worth noting here that the multicellular trichomes of vegetative sterile elements of floral bracts, bracteoles and perianth segments of some chenopods and graminaceous plants are related to salt and heavy metal removal. In some cases, it has been observed that a high concentration of various ions accumulates in the vacuole of bladder trichome terminal cells. There are two types of glandular tri-chomes (salt glands) found by us in Salsola species as against the data presented by others related to the absence of salt glands in chenopods (Carolin 1983). The reason may be that they are not strictly homologous, particularly since both occur in annual Salsola species. We suggest that the different appearance of terminal cells by these two types is due to differences in function connecting both with the accumulation of various ions and /or secretory processes. A comparative morphological study of closely related annual Salsola species from highly contaminated desert soils (Uzbekistan) and unpolluted steppe soils (Europe) shows an increase of succulent bracts/perianth segments consistent with Kyzylkum chenopods, epidermal vesicles were rarely recorded here. The prickles, as single celled hairs with relatively thin cellulose walls and thick cuticles that has been described for some chenopods in some annual chenopods may represent the final stage in the reduction of uniseri-ate hairs (type 3 and 4) according to the classification presented by Carolin (1983). We are inclined to consider various morphological types of hairs described mostly for Salsola species as part of the same transformation series, which probably perform different functions, but little is known about the origin and significance of such transformations, especially when they occur on the same plant.

6 Stomatal Diversity

Stomatal frequency within representatives of genus Salsola varies greatly on different parts of the same leaf/or leaf-like organs and on different leaves, bracts/bracteoles of the same plant and is influenced by environmental conditions. In bracts/bracteoles of Salsola species stomata occur on both sides or mostly or only on one side, usually lower. Stomata also vary in the level of their position on the epidermis. Some are even with the other epidermal cells; others are raised above or sunken below the surface (as in the case of S. lanata, S. turkomanica). The number of stomata per unit area and the positional level of the guard cells with respect to other epidermal cells are so variable that they are of little taxonomic value. The more frequently used taxonomic character is the appearance of the stomata as seen from the surface, especially with reference to the nature and orientation of the neighboring cells). The stomatal counts indicate a great variation in the absolute number per unit area, probably due to differences in variety (species) and ontogenetic stage of leaf-like organs.

A large diversity in the anatomy of assimilatory organs and their photosynthetic pathway has been marked within genus Salsola. Two anatomical types, Salsoloid and Sympegmoid occur in the leaves of species of Salsola (Toderich et al. 2007; Voznesenskaya et al. 2002; Freitag and Stichler 2002; Akhani et al. 2007). In some species with Salsaloid anatomy NAD-ME C4 photosynthesis has been reported, whereas others have NADP-ME C4-subtype (P'yankov et al. 1997, 2001). Plants with Sympegmoid anatomy have C3-like 13C/12C discrimination values (P'yankov et al. 1997, 2000). The variations also occur in structural and biochemical features in cotyledons (P'yankov 1999; Akhani et al. 2007). Two non-Kranz anatomies, Atriplicoid and Salsoloid, are found in Salsola cotyledons (Winter 1981; Butnik et al. 1991; P'yankov et al. 2001), such as cotyledons and leaves may or may not contain a hypodermis. The result is a number of unique combinations of structural and biochemical photosynthetic types in leaves and cotyledons in the species of Salsoleae. So, multiple origins of C4 photosynthesis as described in the families of Poaceae, Cyperaceae, Asteraceae and Zygophyllaceae appear within Chenopodiaceae as well and diversity of photosynthetic types and anatomical structures in the tribe Salsoleae suggests a dynamic pattern of photosynthetic evolution within this single tribe.

From our phenological observations and experimental results, it seems that structural polymorphism of floral organs and sexual reproduction system in some Asian Salsola species are coupled with the diversity of photosynthetic pathways and anatomy of the CO2 assimilative organs. S. arbusculiformis manifests a Sympegmoid leaf and bract anatomy, and non-Kranz bundle sheath cells (Voznesenskaya et al. 2001; Toderich unpublished data). Other species of Section Coccasalsola forming a unique "plant functional group" can be united by Salsoloid (with hypodermis both in leaves and reproductive organs) or a "Crownary-centrical Kranz type of photosynthetic cell arrangement (Voznesenskaya and Gamaley 1986). The anatomy of Salsoloid type of Kranz assimilation tissues is always associated with the C4 syndrome, C4 like 13C/12C carbon discrimination values in leaves, flowers and fruits with a range of 12.0-14.08 (Carolin et al. 1975; Freitag 1997). Such similarity of anatomical and biochemical features is well coordinated with developmental stability of reproductive systems noted by us for S. arbuscula, S. richteri and S. paletzkiana. However, plants of S. arbusculiformis from their natural habitats with Sympegmoid leaf and bract anatomy maintain their 13C/12C, C3/C4 carbon fractionation values in the range from 23.6 to 26.31 throughout their ontogeny, although significant variation was found within plant organs with 2.69% in flowers. This species is also characterized by a set of primitive embryological features such as ana-campylotropous, crassinucellate, bitegmic ovule, autogamy (self pollination /fertilization system), narrow specialization of sexual reproductive system that may be an evidence of lower reproductive plant functional activities leading to the lower level of seed set, seed viability and seed germination. Since C3 is regarded as the primary type of photosynthesis in relation to C4, apparently there is a strong connection between structural floral and fruit traits and their physiological and biochemical activity throughout their ontogeny. The anatomy of bracts in different

Fig. 12.10 a First stage of differentiation of sclerenchyma of S. praecox. b Anatomy and indefinite Kranz bundle sheath cells of S. praecos salsaloid bracts during budding stage. c Cross section of S. praecox bract during flowering; Kranz bundle sheath cells visible. d Cross section of S. praecox bract during fruit maturation

Fig. 12.10 a First stage of differentiation of sclerenchyma of S. praecox. b Anatomy and indefinite Kranz bundle sheath cells of S. praecos salsaloid bracts during budding stage. c Cross section of S. praecox bract during flowering; Kranz bundle sheath cells visible. d Cross section of S. praecox bract during fruit maturation

Asian species of Salsola was studied by us in relation to their photosynthetic activity (Fig. 12.10a-d).

Photosynthetic activity of reproductive organs was insignificant in the budding stage with some increase during flowering process and gradually decreasing during fruit maturation (Figs. 12.10a-d). It was found that S. pestifer, S. praecox and S. paulsenii are similar in photosynthesis types: C4-Sals (-H) both in leaves, cotyledons and bracts. Differences were revealed in the anatomy of bracts. All Asian annual Salsola species of section Salsola subsec. Kali have so-called Salsoloid or 'crown centric' Kranz leaf and bract anatomy (Voznesenskaya and Gamaley 1986). The first features of differentiation of chlorenchyma cells in the bracts and bracteoles are marked at the early stage of pollen sac development, reaching a maximum during blooming stage. Cross sections of perianth in the fruits of many annual Salsola species during maturity also show an insignificant development of chlorenchyma tissue. Similar situation has been described for the species of section Belanthera. In the bracts or fruiting bodies of this type, chlorenchyma is represented by two layers of green cells positioned around the periphery of the organs, the outer layer composed of palisade mesophyll cells and the inner layer composed of palisade mesophyll cells + inner layer of bundle sheath cells. The main vascular bundle with much thick-walled in the centre, surrounded by the water-storage tissue, and only small peripheral bundles have contact with cholernchyma. In fact all species with Salsaloid Kranz anatomy in photosynthetic organs (irrespective of whether these are leaves, stems, cotyledons or bracts) have C4 type photosynthesis (Toderich et al. 2007; P'yankov 1999; P'yankov et al. 2000). However, chlorenchyma of S. ruthenica, consisting of palisade and Kranz cells, is interrupted by longitudinal colenchymatic ridges.

Diversity in the anatomy of fruits reflects the character of adaptive coevolution of woody Salsola taxa and plays a more significant role in the species identification than other elements of floral organs. For instance in S. richteri and S. paletzkiana the adaptive specialization to the xeric-arid conditions proceeds towards the intensification of sclerification of fruiting perianth and increase in the size and number of cell layers of pericarp and even embryo tissues. The presence of pigments in the fruit covers, singular hydrocytic cells, partial myxospermy and development of membranous layer in the spermoderma intensify the defending function against sun radiation. A fully developed embryo and differentiation of its tissues indicates the complete readiness of embryo of Salsola species to the germination. Seed dispersal is manifested by the development of large and wide wings; all elements of fruit cover and embryos of studied species have adaptive value in pigmentation, partial myxospermy, thickenings of external walls, membranous and aleironic layers in the spermoderma, intensification of succulence features as a result of well development of aerial parenchyma, abundance of reserve store nutritional substances, which stimulate the defense mechanism of embryo under extreme desert environments.

The Asian Salsola species of section Arbuscula Coccosalsola section with both C3 and C4 photosynthesis types represent a unique example of the evolutionary convergence of ecological, structural, physiological and biochemical traits. The great range of variation, far more marked in ploidy of genome and fruit structures than in floral and pollen morphology explains the high phenotypic plasticity and good adaptation of S. richteri and S. arbuscula to various geographical and ecological desert habitats. On the other hand S. paletzkiana and S. arbusculiformis are characterized by narrow structural specialization of reproductive organs, partly seeds to germinate only on the sandy or stony gypsumferous soils that, perhaps explains the strict local distribution of this species in the Central Asian Flora (Toderich et al. 2008).

An analysis of the carbon isotope ratio (S13C) of wild Kyzylkum desert species along the salinity gradient revealed significant differences in carbon discrimination between and within C3 and C4 species. Within C3S13C value changes from -30.1%c (Zygophyllum fabago, Zygophyllaceae) to -25.61%c (Tamarix hispida,Tamaricaceae). In general for the C3 plants investigated by us differences in 13C between different species reached 5.49%, and within separate species -3.26% (Alhagi pseudalhagi, Fabaceae). Such changes of carbon discrimination in plants are evidence of change in photosynthetic intensity, as well water use efficiency more than 50%. A 2% difference in the discrimination of C3 species indicates a difference in water-use efficiency of about 30% (Ehleringer and Cooper

1988; Ehleringer et al. 1998). For C4 species the difference in 13C value was not so significant: from — 14.241%c (Kochia prostrata, Chenopodiaceae) to -12.31%e (Suaeda arcuata, Chenopodiaceae).

Stable carbon isotope analysis of different plant communities showed that mean 13 C of C3 species in xerophytes communities was lower, than haloxerophytes and halophytes: —27.39%, —26.67%, h —24.79% . For C4 species in the same community follow results were obtained —12,86% in haloxerophytes, —12.63% xerophytes and —12.16% halophytes. It may be due to various salinity levels of soil, because haloxerophyte and halophyte communities occupy soils with moderate and high level of salinity, whereas xerophytic communities grow on non/light saline soils. A negative effect of soil salinity on carbon isotope ratio of desert plants was observed. In general C3 species are more sensitive to the soil salinity than C4. Salinity, as stress factor decreases the transpiration and photosyn-thetic intensity, which leads to a decrease in the rate of biomass accumulation of plants.

7 Conclusion

In conclusion it can be said that desert plants as autotrophic sessile organisms are continuously facing changing and unpredictable environments as well as micro-environmental problems to solve the problems within their organs and cell types which continuously face changing supplies of nutrient ions, sugars, amino acids, gases, light and water. Some major external environmental problems that plants must solve are:

- their biophysical soil and air environments are continuously changing, far beyond normal daily environmental changes;

- their biological environments (microbes, herbivores, and others) change constantly;

- human's particularly move and destroy plants and add both beneficial materials and toxic pollutants to their environments

In the Kyzylkum Desert some plants;characterized as metallohalophytes by us; grow well in either natural or contaminated soils containing salts and metals (Toderich et al. 2004a, b, 2005a, b, 2006). The flora in this desert contains only a restricted number of species capable of removing metal/salts from their habitats. These species can survive and reproduce under these contaminated environments. Some successful species in such habitats produce large quantities of small, easily dispersed seeds, hence facilitating colonization. It is clear from the biochemical and physiological studies that plants have multiple often redundant pathways and mechanisms to accomplish the same function or goal. These genetically built-in mechanisms for redundancy in numerous plant functions act as fail-safe mechanisms. Redundancy apparently gives sessile plants 2 major advantages;

1. their normal developmental ability to form diverse functions in different types of organs, tissues and cells,

2. a very powerful means to adapt the functions of these structures to cope with whatever happens in their biophysical and biological environments (Black 1993; Black etal. 1995).

As external environmental CO2 levels vary, the internal CO2 levels in green photosynthetic tissues can be modified to provide this essential nutrient (Toderich et al. 2007).

There are several morphological and anatomical features met within desert plants under natural saline and contaminated environments but most important ones are salt-secretary trichomes and salt glands. These resemble functionally and are associated with the secretion of ions using morphological characters, mainly related to epidermal structures (by SEM analysis). Desert species are developing different mechanisms of adaptation to stress; species with salt-producing trichomes/hairs and salt-accumulating (with specific salt/storage cells) plants. This is an indication that different mechanisms and strategies for the sequestration and regulation of the salt ion concentration in the plant tissues are operated in the stem and leaf succulent halophytes and in the recreto-and pseudohalophytes of the Kyzylkum flora. The existence of great diversity in photosynthetic pathways ofAsiatic Salsola species, as well as anatomy and biochermical features in the CO2 assimilation organs is evidence related to plant growth, survival, and reproduction in such desert plants (Butnik et al. 2001a, b; P'yankov et al. 2001, 2002).

Various morphological types of hairs described mostly for Salsola species as part of the same transformation series probably perform different functions. However little is known about the origin and significance of such kind of transformations, especially when they occur on the same plant.

Increasing of sclerification, availability of pigments and tracheids like cells holding moisture, abundance of crystals in the fruit tepals, tissues also promote the protection of embryo from unfavourable conditions (Butnik et al. 2001a, b; Toderich et al. 2008). Some highly adapted metallohalophytes in nature develop a cellular mechanism to partition toxic salts into vacuoles or to exclude salt at the root zone so it does not affect cell metabolism and division, i.e., a high concentration of various ions can accumulate in the vacuoles of bladder-trichome terminal cells which are frequently developed on the adaxial surface of epidermal cells of leaves or bract/bracteoles

The prominent levels of sclerification of perianth segments combined with thickening of pericarp and spermoderma epidermis bearing papillae-shaped pro-trubarences (Salsola paulsenii) are related to the defending of embryo against entrance of toxic elements. Diversities in sexual reproduction mechanisms and CO2 fixation pathways, for tree-like Salsola species, also are important factors regarding reproduction and survival under saline and technogenic contaminated desert environments.

Most essential plant nutrients come from soil-plant interactions via root and microbial contacts; simultaneously essential nutrient uptake must cope with the presence of any toxicants and non-essential elements in soils. The roles of fungi, bacteria, and other organisms as they interact with plants are crucial. Biological lipid bilayer membranes are essentially impermeable to ions, sugars, and polar molecules; hence channels, pumps, diffusion, solution, and mass flow are used to cross biological membranes. The uptake of mineral ions from soils by plant roots occurs through protein-built channels in a biphasic fashion, first with a strong high affinity active carrier mechanism, followed by a slower diffusion uptake. Such active transport channels and pumps are powered, usually by ATP, and may involve an active co-transport with other ions or an exchange with others ions.

For bioremediation purposes there should be interest in the species which consistently have a metal/salt removal potential. Since several "hyperaccumulators" are characterized by small biomass production, the use of selected metallohalophyte species as phytoremediators capable of accumulating high amounts of toxic ions should be considered. Halophytes and simultaneously metal tolerant arid/semiarid plants may be used for phytoremediation of areas contaminated with toxic salts and heavy metals. However, future work is needed to:

- Select optimal genotypes from Kyzylkum desert flora and to initiate a program of its seed multiplication.

- Determine the mechanisms of their hyperaccumulation and hypertolerance.

- Isolate the genes involved.

It may then be possible to genetically engineer these traits into higher biomass forms and develop more efficient heavy metal phytoextraction processes. Several authors have pointed out that heavy metal hyperaccumulators could prove economically useful as an efficient method for cleaning the soils (Leblane et al. 1999; Escarre et al. 2000; Chaney et al. 2007). Significant progress has been made in recent years in developing native or genetically modified plants for the remediation of contaminated sites (Meagher et al. 2000). The study of chemical compounds (origin, localization etc.) for Asian desert plants are of great interest because they are often specific to a particular plant species or genus and must therefore have been designed to serve a particular protective function. In the case of salt remediation the timing of salt excretion within plant organs is of critical importance, not only for our understanding of the cellular mechanism involved, but also because salt/toxin accumulation could interfere with health problems of other living beings.

The stable recovery of ecosystem functions can be considered best from the viewpoint of development over time. Phytoremediation technology is considered a potentially valuable technique for dealing with heavy metals, which are typically the most difficult pollutants to remove from soils. The use of metallohalophytes from the Central Asian flora to reclaim soils could represent both a practical and economically viable strategy. Even though the scientific technology for molecu-larly transforming plants is very well established, unfortunately plants that are well adapted to desert environments have not yet been transformed. Plant transformation knowledge needs to be applied immediately to the special needs of desert-adapted plants in Central Asia.

The cultivation of halophytes (C3 and C4 plants) can limit long-distance salt spreading and improve the vitality and growth conditions for local species, when cultivated together. Since stress conditions frequently trigger defense mechanisms based on the production of specific biological active metabolites of pharmaceutical or industrial importance, halo-metallophytes of the South part of Aral Sea Basin could constitute a valuable source of cash compounds. These characteristics may offer a new and valuable source of income to local populations.


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