Morphological and anatomical contributions to the taxonomical identification of two <Emphasis Type="Italic">Ornithogalum</Emphasis> taxa (<Emphasis Type="Italic">O. nutans and O. boucheanum</Emphasis>) from Flora of Turkey

In this study, morphological and anatomical features of Ornithogalum nutans and O. boucheanum, two relative and morphologically similar species growing in European Turkey, were investigated. These species showed some important anatomical differences with regard to leaf anatomy while they have identical features in stem. The stem anatomy of these two species displays the common properties of monocotyledons. The mesophyll is unifacial and contains monotypic chlorenchyma cells in the leaf of O. nutans. It has no lacunae. The mesophyll in O. boucheanum is equifacial and it has lacunae. This anatomical distinction may be useful for the identification of these similar-looking species.     

Restionaceae seedlings: Morphology,anatomy and systematic implications

Restionaceae differ from most monocot families in having both epigeal and hypogeal germination. The green cotyledons associated with epigeal germination have a central vascular strand as found in most epigeal monocotyledons. In some genera the cotyledon may have a hairpin‐like structure, also described for Anthericaceae. The cotyledon of the hypogeal seedlings is short, without green pigment and largely remains embedded in the seed coat. Hypogeal germination is correlated with large, woody, indehiscent, frequently myrmecochorous nuts, while epigeal germination is found in species with smaller indehiscent nutlets or seeds, dispersed in a variety of ways. The primitive condition is most likely epigeal germination. In hypogeal seedlings of some African and Australian taxa an epicotyledonary rhizome is found between the primary root and the first leaves. Seedlings of African Restionaceae frequently have elongated culm internodes, whereas in the Australian species studied, internodes are very short, resulting in a cluster of seedling leaves. The leaf blades, which in most species are only found on the seedlings, are very simple anatomically. However, they appear to be unifacial, similar to the leaf blades of Anarthria (Anarthriaceae). The anatomical specialisations in the blades mirror those recorded for the culm anatomy. These observations are consistent with the hypothesis that Centrolepidaceae may be neotonous Restionaceae. They also corroborate the morphology of the African Restionaceae, and the presently accepted phylogeny of the African genera of Restionaceae.     

Evolutionary and developmental studies of unifacial leaves in monocots: Juncus as a model system

An important objective in evolutionary developmental biology is to understand the molecular genetic mechanisms that have given rise to morphological diversity. Leaves in angiosperms generally develop as a flattened structure with clear adaxial–abaxial polarity. In monocots, however, a unifacial leaf has evolved in a number of divergent species, in which leaf blades consist of only the abaxial identity. The mechanism of unifacial leaf development has long been a matter of debate for comparative morphologists. However, the underlying molecular genetic mechanism remains unknown. Unifacial leaves would be useful materials for developmental studies of leaf-polarity specification. Moreover, these leaves offer unique opportunities to investigate important phenomena in evolutionary biology, such as repeated evolution or convergent evolution of similar morphological traits. Here we describe the potential of unifacial leaves for evolutionary developmental studies and present our recent approaches to understanding the mechanisms of unifacial leaf development and evolution using Juncus as a model system.     

单子叶植物科级分类阶元形态性状分支分析

基于93个形态形状,采用13个被子植物基部类群做为外类群,对49个单子叶植物科级分类阶元进行了分支系统学分析。经过简约性分析,得到了1684棵同等最大简约分支树。严格一致树的分支结构图表明：1）古草本类植物和单子叶植物是姐妹群关系;2）具有网状脉的类群,薯蓣科,菝葜科,百部科是单子叶植物的最基部类群。由于性状状态间存在着较多的平行和逆转进化,这在一定程度上影响了系统发育重建的准确性;所选择的性状状态之间的演化很可能是平行的、多次的或者是特化的状态,因此这样复杂的演化关系的探索关键在于找到一些能确切反映其系统演化关系的形态性状。目前很难通过简约化的形态分支分析来解开整个单子叶植物的起源和演化之谜。为了避开对系统学分析造成干扰的误导性状,形态数据结合DNA序列分析很可能是必需的。     

A Role for Auxin in Triggering Lamina Outgrowth of Unifacial Leaves

A common morphological feature of typical angiosperms is the patterning of lateral organs along primary axes of asymmetry—a proximodistal, a mediolateral, and an adaxial–abaxial axis. Angiosperm leaves usually have distinct adaxial–abaxial identity, which is required for the development of a flat shape. By contrast, many unifacial leaves, consisting of only the abaxial side, show a flattened morphology. This implicates a unique mechanism that allows leaf flattening independent of adaxial–abaxial identity. In this study, we report a role for auxin in outgrowth of unifacial leaves. In two closely related unifacial-leaved species of Juncaceae, Juncus prismatocarpus with flattened leaves, and Juncus wallichianus with transversally radialized leaves, the auxin-responsive gene GLYCOSIDE HYDROLASE3 displayed spatially different expression patterns within leaf primordia. Treatment of J. prismatocarpus seedlings with exogenous auxin or auxin transport inhibitors, which disturb endogenous auxin distribution, eliminated leaf flatness, resulting in a transversally radialized morphology. These treatments did not affect the radialized morphology of leaves of J. wallichianus. Moreover, elimination of leaf flatness by these treatments accompanied dysregulated expression of genetic factors needed to specify the leaf central-marginal polarity in J. prismatocarpus. The findings imply that lamina outgrowth of unifacial leaves relies on proper placement of auxin, which might induce initial leaf flattening and subsequently act to specify leaf polarity, promoting further flattening growth of leaves.

Lamina outgrowth of unifacial leaves, which lack adaxial identity, relies on proper localization of auxin, which might induce initial leaf flattening and subsequently act to specify leaf polarity, promoting further flattening growth of leaves.     

Shoot apex,leaf development and unifacial tip inAgave wightii drumm. et prain (agavaceae)

The shoot apex has one tunica layer enclosing a mass of corpus which is differentiated cytohistologically into central mother cell zone, flank zone, rib zone and a ‘cambium-like’ zone. Occurrence of ‘cambium-like’ zone during minimal phase is considered as an expression of nodal region. Agave wightii shows spirodistichous arrangement of leaves which have an expanded photosynthetic surface with a reduced unifacial tip. Leaves are initiated by periclinal divisions in the second layer. Vertical growth in the leaves is by subapical initials and lateral growth is by marginal and submarginal initials in their early stages of development. The unifacial tip is formed by the extension of adaxial meristematic activity. The derivatives thus formed are pushed to the abaxial side of the primordiuj. Hence the unifacial part of the leaf is regarded as equivalent to a phyllode.     

Lateral meristems and stem thickening growth in monocotyledons

Although monocotyledons lack a vascular cambium of the type found in dicotyledons and conifers, lateral meristems still play an important role in the establishment of their growth habits. The presence near the shoot apex of a primary thickening meristem (PTM), which is probably plesiomorphic in monocotyledons, predisposes evolution into the many pachycaul forms. A PTM occurs in virtually all monocotyledons, whereas the secondary thickening meristem (STM), which is morphologically similar, is limited to a few genera of Liliiflorae. these records are reviewed in a systematic context. To a greater or lesser extent in different taxa, the PTM is responsible for primary stem thickening, adventitious root production, and formation of linkages between stem, root and leaf vasculature. The STM largely contributes to the body of the stem. The sometimes obscure distinction between the two meristems, and their relationship with other stem meristems are discussed. For systematic purposes stem thickening in monocotyledons is separated into two characters: diffuse growth (as in palms), and growth by means of lateral meristems. The three states of the second character are represented by the first three of Mangin’s (1882) four categories (two herbaceous, the third arborescent): (1) The lateral meristem is limited in extent, and ceases activity after root formation. (2) It remains active for a limited period after cessation of root formation, contributing to the plant body. (3) It remains active throughout the life of the plant, contributing the bulk of the plant body.     

Systematic root anatomy of Asparagales and other monocotyledons

Root anatomy of several taxa of Asparagales and some taxa formerly included in Asparagales is described in a systematic context together with a literature review. The presence of a dimorphic outer layer with long and short cells is widespread in monocotyledons, indicating that it originated early in the monocot lineage, but whereas this layer is rhizodermal in most monocotyledons, in Asparagales and Araceae it is usually hypodermal. There may be a correlation between the presence of a velamen or a persistent rhizodermis in many Asparagales and Araceae and the presence of a dimorphic hypodermal layer. Many other root anatomical characters, such as the presence of vascular bundles in the central pith and a multi-layered sclerenchymatous cylinder, are probably xeromorphic and developed convergently.     

LEAF DEVELOPMENT IN SENECIO ROWLEYANUS (COMPOSITAE)

Prolonged apical growth of the leaf primordium and the presence of distinct marginal meristems do not occur in Senecio rowleyanus. Intercalary cell divisions accompanied by radial expansion of derivatives from an adaxial meristem account for the spherical shape of the leaf. The “window” in the lamina marks the position of the adaxial meristem and precludes interpretation of the leaf as being unifacial. Stomata are mesoperigenous and anomocytic in type. Schizogenous secretory canals occur in both the leaf and the stem, and their association with vascular bundles is discussed. The anatomy of the leaf is interpreted in terms of xeromorphy.     

南昌市不同植物类群叶片氮磷浓度及其化学计量比

对南昌大学前湖校区89种主要植物叶片的N、P浓度及其化学计量比进行了研究,结果表明:乔灌、常绿、针叶、种子、裸子和单子叶植物类群的N浓度分别低于相对应的草本、落叶、阔叶、蕨类、被子和双子叶植物类群,而C3和C4植物差异不显著;乔灌、常绿和裸子植物类群的P浓度含量分别低于相对应的草本、落叶和被子植物类群,而针叶和阔叶、蕨类和种子、单子叶和双子叶、C3和C4植物类群间差异不显著;乔木、阔叶、被子和双子叶植物类群叶片N/P分别高于相对应的灌草、针叶、裸子和单子叶植物类群,而常绿和落叶、蕨类和种子、C3和C4植物类群之间差异不显著.可见,不同类型植物对N和P的吸收利用存在差异,且对不同养分供应采取不同的适应对策.结合研究区土壤养分现状,建议优先选择常绿、针叶、裸子和单子叶植物类群作为城市园林植物.     

COMPARATIVE FOLIAR HISTOGENESIS IN ACORUS CALAMUS AND ITS BEARING ON THE PHYLLODE THEORY OF MONOCOTYLEDONOUS LEAVES

A comparative histogenetic investigation of the unifacial foliage leaves of Acorus calamus L. (Araceae; Pothoideae) was initiated for the purposes of: (1) re-evaluating the previous sympodial interpretation of unifacial leaf development; (2) comparing the mode of histogenesis with that of the phyllode of Acacia in a re-examination of the phyllode theory of monocotyledonous leaves; and (3) specifying the histogenetic mechanisms responsible for morphological divergence of the leaf of Acorus from dorsiventral leaves of other Araceae. Leaves in Acorus are initiated in an orthodistichous phyllotaxis from alternate positions on the bilaterally symmetrical apical meristem. During each plastochron the shoot apex proceeds through a regular rhythm of expansion and reduction related to leaf and axillary meristem initiation and regeneration. The shoot apex has a three- to four-layered tunica and subjacent corpus with a distinctive cytohistological zonation evident to varying degrees during all phases of the plastochron. Leaf initiation is by periclinal division in the second through fourth layers of the meristem. Following inception early growth of the leaf primordium is erect, involving apical and intercalary growth in length as well as marginal growth in circumference in the sheathing leaf base. Early maturation of the leaf apex into an attenuated tip marks the end of apical growth, and subsequent growth in length is largely basal and intercalary. Marked radial growth is evident early in development and initially is mediated by a very active adaxial meristem; the median flattening of this leaf is related to accentuated activity of this meristematic zone. Differentiation of the secondary midrib begins along the center of the leaf axis and proceeds in an acropetal direction. Correlated with this centralized zone of tissue specialization is the first appearance of procambium in the center of the leaf axis. Subsequent radial expansion of the flattened upper leaf zone is bidirectional, proceeding by intercalary meristematic activity at both sides of the central midrib. Procambial differentiation is continuous and acropetal, and provascular strands are initiated in pairs in both sides of the primordium from derivatives of intercalary meristems in the abaxial and adaxial wings of the leaf. Comparative investigation of foliar histogenesis in different populations of Acorus from Wisconsin and Iowa reveals different degrees of apical and adaxial meristematic activity in primordia of these two collections: leaves with marked adaxial growth exhibit delayed and reduced expression of apical growth, whereas primordia with marked apical growth show, correspondingly, reduced adaxial meristematic activity at equivalent stages of development. Such variations in leaf histogenesis are correlated with marked differences in adult leaf anatomy in the respective populations and explain the reasons for the sympodial interpretation of leaf morphogenesis in Acorus and unifacial organs of other genera by previous investigators. It is concluded that leaf development in Acorus resembles that of the Acacia phyllode, thereby confirming from a developmental viewpoint the homology of these organs. Comparison of development with leaves of other Araceae indicates that the modified form of the leaf of Acorus originates through the accentuation of adaxial and abaxial meristematic activity which is expressed only slightly in the more conventional dorsiventral leaf types in the family.     

Pollen and anther characters in monocot systematics

Pollen and anther characters are potentially informative in higher-level systematics of monocotyledons. Several characters of monocot pollen and anthers (tapetum type, microsporogenesis type and inaperturate pollen) are reviewed here in relation to recent phylogenetic concepts of the group, and new data are presented for some critical taxa. The first-branching monocotyledon, Acorus , has a secretory tapetum but most other early branching taxa (i.e., most Alismatales, except Tofieldia ) are plasmodial. The lilioid orders, Pandanales, Dioscoreales, Liliales and Asparagales are almost uniformly secretory. The tapetum is more diverse within the commelinoid clade. Successive microsporogenesis predominates in the monocotyledons although the simultaneous type is of systematic significance within some orders, such as Dioscoreales,Asparagales and Poales. Inaperturate pollen (either "functionally monoaperturate" or "omniaperturate") occurs in every major monocot group. It predominates in Alismatales and Zingiberales, and is a synapomorphy for some Liliales and Asparagales.     

New costapalmate palm leaves from the Oligocene Ningming Formation of Guangxi,China, and their biogeographic and palaeoclimatic implications

New palm leaves from the Oligocene Ningming Formation are placed into the morphogenus Sabalites because of their costapalmate leaf shape. Four taxa are described on the basis of leaf compressions with cuticular structure. S. guanxiensis sp. nov. is characterised by hypostomatic leaf blades with a stout costa and a symmetrical base. Sabalites cf. asymmetricus has amphistomatic leaf blades with a long, delicate costa and an asymmetrical base. Sabalites sp. 1 has amphistomatic leaf blades with a long, massive costa and wide segments. Sabalites sp. 2 is characterized by hypostomatic leaf blades with a prominent costa and an asymmetrical base. The four new palm taxa expand our understanding of the floristic elements and features of the Oligocene Ningming flora. Together with the other three palm taxa that were previously reported from the Ningming Formation, our material indicates that the Oligocene Ningming flora had a rich diversity of costapalmate palms. The relatively high species diversity of palms and other plants corroborate that the Oligocene Ningming flora represents a warm and humid climate.     

LEAF-OPPOSED BUDS IN MUSA: THEIR DEVELOPMENT AND A COMPARISON WITH ALLIED MONOCOTYLEDONS

A single, lateral, vegetative bud which is positioned 180° from the axil of a leaf is a generic feature of Musa (Musaceae). Such leaf-opposed buds occur in all ten species and five cultivars examined, representing all four sections of the genus and all groups of cultivated bananas and plantains. The bud arises relatively late and is first visible as a vascular-free “clear zone” in the axis directly below the future bud meristem site. It is first associated with the fifth or sixth leaf primordium from the apex. A defined superficial meristem develops on the stem directly above the insertion of the leaf margins one or more plastochrons later. Normal, basically axillary, vegetative buds occur in the closely related genera: Orchidantha (Lowiaceae), Heliconia (Heliconiaceae), Strelitzia, and Ravenala (Strelitziaceae). These buds arise in the axil of the first to the third leaf primordium in a manner similar to most other monocotyledons. Axillary vegetative buds also occur in the remaining families of the Zingiberales: Cannaceae, Costaceae, Marantaceae, and Zingiberaceae.     

Do plant mites commonly prefer the underside of leaves?

The adaxial (upper) and abaxial (lower) surfaces of a plant leaf provide heterogeneous habitats for small arthropods with different environmental conditions, such as light, humidity, and surface morphology. As for plant mites, some agricultural pest species and their natural enemies have been observed to favor the abaxial leaf surface, which is considered an adaptation to avoid rain or solar ultraviolet radiation. However, whether such a preference for the leaf underside is a common behavioral trait in mites on wild vegetation remains unknown. The authors conducted a 2-year survey on the foliar mite assemblage found on Viburnum erosum var. punctatum, a deciduous shrub on which several mite taxa occur throughout the seasons, and 14 sympatric tree or shrub species in secondary broadleaf-forest sites in Kyoto, west–central Japan. We compared adaxial–abaxial surface distributions of mites among mite taxa, seasons, and morphology of host leaves (presence/absence of hairs and domatia). On V. erosum var. punctatum, seven of 11 distinguished mite taxa were significantly distributed in favor of abaxial leaf surfaces and the trend was seasonally stable, except for Eriophyoidea. Mite assemblages on 15 plant species were significantly biased towards the abaxial leaf surfaces, regardless of surface morphology. Our data suggest that many mite taxa commonly prefer to stay on abaxial leaf surfaces in wild vegetation. Oribatida displayed a relatively neutral distribution, and in Tenuipalpidae, the ratio of eggs collected from the adaxial versus the abaxial side was significantly higher than the ratio of the motile individuals, implying that some mite taxa exploit adaxial leaf surfaces as habitat.     

THE PRIMARY THICKENING MERISTEM: DEFINITION AND FUNCTION IN MONOCOTYLEDONS

The PTM should be defined as a diffuse primary meristem which decreases in cross-sectional extent (i.e., becomes a thinner-walled cylinder) in a basipetal direction. It is associated with extensive anticlinal cell files and consists of cell initials that divide predominantly in periclinal planes. This meristem occurs typically in monocotyledons, especially those with thick, compact stems in species with rosette shoot axes. The PTM is also associated with a wide crown, so that the apical meristem is either slightly above the level of youngest leaf primordia, at approximately the same level as the leaf primordia, or distinctly sunken below surrounding stem tissue and the youngest leaf primordia. The location is dependent on the extent of primary thickening growth occurring in a particular species. A meristem associated with primary thickening of other plant groups should not be called a primary thickening meristem unless all of the above characteristics are shown to be associated with the meristem being examined. The primary thickening meristem is responsible for primary thickening of a stem axis. Its ontogenetic relationship with the STM needs further investigation. Extensive primary stem thickening has been observed in non-monocotyledons (ferns, lycopods, cycads, and dictyledons). Some of these organisms appear to undergo primary thickening from a PTM in a similar process as that which occurs in monocotyledons. Further research is necessary to establish the mechanisms of primary thickening in these cases.     

Sieve-element plastids and evolution of monocotyledons, with emphasis on Melanthiaceae sensu lato and Aristolochiaceae-Asaroideae, a putative dicotyledon sister group

Monocotyledons are distinguishable from dicotyledons by their subtype P2 sieve-element plastids containing cuneate protein crystals, a synapomorphic character uniformly present from basal groups through Lilioids to Commelinoids. The dicotyledon generaAsarum andSaruma (Aristolochiaceae-Asaroideae) are the only other taxa with cuneate crystals, but their sieveelement plastids include an additional large polygonal crystal, as is typical of many eumagnoliids. New investigations in Melanthiaceae s.l. revealed the same pattern (polygonal plus cuneate crystals) in the sieve-element plastids ofJaponolirion osense (Japonoliriaceae/Petrosaviaceae), ofHarperocallis flava, Pleea tenuifolia, andTofleldia (all: Tofieldiaceae). InNarthecium ossifragum a large crystal, present in addition to cuneate ones, usually breaks up into several small crystals, whereas inAletris glabra andLophiola americana (Nartheciaceae) and in all of the 15 species studied and belonging to Melanthiaceae s.str. only cuneate crystals are found. Highresolution TEM pictures reveal a crystal substructure that is densely packed in both cuneate and polygonal forms, but in Tofieldiaceae the polygonal crystals stain less densely, probably as a result of the slightly wider spacing of their subunits. The small crystals ofNarthecium are “loose”; that is, much more widely spaced. Such “loose” crystals are commonly found in sieve-element plastids of Velloziaceae, present there in addition to angular crystals, and together with cuneate crystals in a few Lilioids and many taxa of Poales (Commelinoids). Ontogenetic studies of the sieve elements ofSaruma, Aristolochia, and several monocotyledons have shown that in their plastids cuneate crystals develop very early and independent from a polygonal one present in some taxa. Therefore, a conceivable particulation of polygonal into cuneate crystals is excluded. Consequently, mutations of some monocotyledons that contain a lone, large, polygonal crystal in their sieve-element plastids are explained as the result of a complex genetic block. The total result of all studies in sieve-element plastids suggests thatJaponolirion and Tofieldiaceae are the most basal monocotyledons and that Aristolochiaceae are their dicotyledon sister group.     

The diversity of stomatal development in Orchidaceae subfamily Orchidoideae

The development of leaf stomata in species of Orchidoideae sensu Garay is agenous or hemimesogenous with a single mesogene cell. Both kinds of development occur in all 26 species studied and are found in a characteristic proportion which may differ markedly even between related species.
Leaf and stem stomata develop similarly but surrounding cells may divide obliquely in the latter, so that perigene cells are formed. At maturity, mesogene and perigene cells resemble other epidermal cells, the stomata being anomocytic.
In monocotyledons, the hemimesogenous development of stomata is previously known only from five orchid genera of the neottioid tribe Cranichideae sensu Dressler. This kind of stomatal development in monocotyledons is documented by micrographs for the first time. Mesogene and perigene cells are recorded for the first time in the Orchidoideae. The diversity of types of stomatal development in this group is emphasized.     

Leaf adaxial-abaxial polarity specification and lamina outgrowth: evolution and development

A key innovation in leaf evolution is the acquisition of a flat lamina with adaxial-abaxial polarity, which optimizes the primary function of photosynthesis. The developmental mechanism behind leaf adaxial-abaxial polarity specification and flat lamina formation has long been of interest to biologists. Surgical and genetic studies proposed a conceptual model wherein a signal derived from the shoot apical meristem is necessary for adaxial-abaxial polarity specification, and subsequent lamina outgrowth is promoted at the juxtaposition of adaxial and abaxial identities. Several distinct regulators involved in leaf adaxial-abaxial polarity specification and lamina outgrowth have been identified. Analyses of these genes demonstrated that the mutual antagonistic interactions between adaxial and abaxial determinants establish polarity and define the boundary between two domains, along which lamina outgrowth regulators function. Evolutionary developmental studies on diverse leaf forms of angiosperms proposed that alteration to the adaxial-abaxial patterning system can be a major driving force in the generation of diverse leaf forms, as represented by 'unifacial leaves', in which leaf blades have only the abaxial identity. Interestingly, unifacial leaf blades become flattened, in spite of the lack of adaxial-abaxial juxtaposition. Modification of the adaxial-abaxial patterning system is also utilized to generate complex organ morphologies, such as stamens. In this review, we summarize recent advances in the genetic mechanisms underlying leaf adaxial-abaxial polarity specification and lamina outgrowth, with emphasis on the genetic basis of the evolution and diversification of leaves.