Correlations between leaf structural traits and the densities of herbivorous insect guilds

The functional composition of herbivorous insect assemblages was correlated with aspects of new and mature leaf surface features, anatomy and morphology across 18 co‐occurring plant species. Multivariate analyses of insects and leaf traits revealed that the functional composition of the herbivore assemblage was more strongly correlated with leaf structural traits than with leaf constituents. Leaf traits were more strongly correlated with the functional composition of the herbivore assemblage than with its taxonomic composition. Densities of sessile phloem feeders, ­rostrum chewers, and all herbivores were significantly negatively correlated with specific leaf weight, lamina and cuticle thickness, vascular tissue depth and stomate length, and were significantly positively correlated with stomate density. External chewer densities were significantly negatively correlated with percent lignified vein area, and significantly positively correlated with leaf surface area and the distance between lignified tissues. Spine‐like leaves were associated with significantly lower densities of sessile phloem feeders, external chewers and all herbivores compared to kite leaves (kite leaves are comprised of unfortified leaf tissue supported by a framework of vascular tissue). The presence of a thickened leaf hypodermis was associated with significantly lower densities of external chewers and rostrum chewers, while midrib protection was associated with significantly lower densities of external chewers. Leaf structural traits may not be the proximal factors influencing herbivorous insects, as leaf structural traits are correlated with many other plant traits such as photosynthetic rate, relative growth rate and leaf life‐span. Nonetheless, these results indicate that certain leaf structural traits may potentially be used to predict the functional structure of herbivorous insect assemblages. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 77 , 43–65.     

Leaf and stem anatomy of Vochysiaceae in relation to subfamilial and suprafamilial systematics

Several leaf anatomical features are potentially systematically informative within both the family Vochysiaceae and the order Myrtales, notably tracheoidal idioblasts, mucilage cells and secretory canals. Tracheoids with spiral wall thickenings are present in the mesophyll of most species of Vochysia , and also occur in several other families of Myrtales. Mucilage cells are common in the leaf epidermis in some Vochysiaceae. Secretory ducts are present in the midrib in Salvertia and Vochysia , which are apparently closely related, although Salvertia also shares some leaf anatomical characters with Qualea and Callisthene . Anatomical data do not support the segregation of Ruizterania from Qualea ; leaves of R. albiflora leaves are very similar to those of Q. paraensis in venation pattern, and leaf and stem anatomy. Different venation patterns are characteristic of sections within the genus Qualea , but within the large genus Vochysia , leaf anatomy is variable even within a subsection. Amongst other Myrtales, leaf anatomy of Vochysiaceae most closely resembles that of Combretaceae and Onagraceae. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 138 , 339–364.     

The systematic importance of anatomical data in Gagea (Liliaceae) from the Flora Iranica area

Anatomical features of basal leaves, pedicels and tepals of 22 species of Gagea belonging to four sections (Platyspermum, Plecostigma, Gagea and Didymobulbos) are investigated. Anatomical characters are mapped onto a molecular phylogenetic tree and their evolution is evaluated. The systematic importance of the anatomical characters is discussed. Anatomical characters are more systematically useful in the Irano‐Turanian taxa than in Euro‐Siberian taxa. The presence of collenchyma and/or sclerenchyma and the pentagonal outline of the transverse section of the basal leaf is found in Irano‐Turanian taxa of section Platyspermum and is mostly absent in Euro‐Siberian taxa. A diagnostic key based on combined anatomical characters is provided. The level of variation in anatomical characters is greater in the basal leaf than the pedicel and lowest in the tepals. Convergent evolution in anatomical characters is associated with ecological shifts between sunny, open, dry habitats and closed, humid habitats. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 155–177.     

The leaf internal morphology and ultrastructure of Zostera muelleri irmisch ex aschers. (Zosteraceae): a comparative study of the intertidal and subtidal forms

The leaf anatomy, histochemistry and ultrastructure of the intertidal and subtidal seagrass Zostera muelleri Irmish ex Aschers. from Westernport Bay, Victoria were studied. Unusual anatomical and ultrastructural features are compared with other seagrasses and their functional significance is assessed. Subcuticular cavities are present in the young blade, but not observed in the older blade nor young and old leaf sheath. Wall ingrowths occur in the blade epidermal cells particularly on the inner tangential walls and the lower portions of the radial walls. Plasmodesmata are present between adjacent epidermal cells and between the epidermal and mesophyll cells, suggesting that solutes could transfer between these tissues both symplastically and apoplastically. Each leaf has three longitudinally aligned vascular bundles, each of which comprises a single xylem element isolated from the phloem tissue. The phloem consists of nacreous-walled sieve elements accompanied by phloem parenchyma cells which also process wall ingrowths. The xylem walls are completely hydrolysed and the middle lamella borders directly on the xylem lumen. Leaves have prominent air lacunae bisected transversely by septa at regular intervals along their length. Each septum consists of a file of small parenchyma cells with wall protuberances projecting into intercellular space. There are no major structural differences between the subtidal and intertidal plants, but the former have larger leaves and more leaves per shoot than the latter. In addition, a network of unusual reticulated fungal hyphae is present in the leaf intercellular spaces of the subtidal form and this network may facilitate solute transfer in these plants.     

The uniqueness of palms

Palms build tall trees entirely by primary growth in a way that limits their growth habit, but not their capacity for continued stem development. They achieve massive primary stature because of distinctive features of leaf development, stem vasculature and anatomical properties. They exhibit several record features of leaf and seed, and inflorescence size and leaves of great complexity. A marked ability to generate new roots allows them to be transplanted easily. As climbing plants they develop the longest unrooted stems in which there are, paradoxically, anomalous features of vascular construction compared with tree palms. It is here claimed that they are the world's longest lived trees because stem cells of several kinds remain active in differentiated tissues throughout the life of the palm. Absence of physiological dormancy may be related to this property, together with inability to withstand freezing temperatures that would cause irreversible cavitation of tracheary elements. This largely restricts them to the tropics, for which they are emblematic organisms. In these biological features palms are indeed unique organisms.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 5–14.     

Systematic and comparative anatomy of Cymbidieae (Orchidaceae)

Cymbideae comprise an assemblage of 28 genera nearly all of which are represented in this study. Their anatomy is relatively homogenous with the exception of Govenia , in which roots lack velamen and pseudobulb vascular bundles lack sclerenchyma, conditions that do not obtain in other genera. Marginal fibre bundles in leaves of Grammatophyllum and Porphyroglottis consist of clusters of thicker-walled, narrower, epidermis-facing fibres as well as thinner-walled, wider, mesophyll-facing fibres. This feature also occurs in some species of Maxillaria . Baculate tilosomes appear in the roots of a majority of genera in Cymbidieae, as they do in species of Maxillaria , confirming DNA analyses showing a close relationship between tribes Cymbidieae and Maxillarieae. Govenia is singled out both on anatomical and molecular grounds as being ill-placed in Cymbidieae. Cladistic analysis produces only a few tentative hypotheses of phylogenetic relationships among the 28 genera, showing that anatomical characters are of limited value in assessing affinities within this tribe. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 1–27.     

The marginal band, a new structural component of the developing angiosperm leaf

The marginal band, a newly described structure of the dicot leaf, plays an essential role in the formation of this organ. It arises along the two edges of a leaf as several files of adaxial epidermal cells during the peg stage of ontogeny. Functionally, it appears to serve as the prepattern for a number of secondary features including (i) the marginal growth meristem present in all leaves as seen in mountain laurel, (ii) rows of lobes and spines as in Salvia and Ilex , (iii) pigmentation as in some cultivars of Hydrangea , (iv) propagation as in the plantlets of Kalanchoe pinnata , (v) marginal ridges for support in a few varieties of Viburnum and Ilex , and (vi) marginal necrosis as in Strelitzia for blade partitioning.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 21–28.     

Comparative vegetative anatomy and systematics of Cymbidium (Cymbidieae: Orchidaceae)

The genus Cymbidium (Orchidaceae) exhibits distinctive ecological diversification and occurs in terrestrial, epiphytic, and lithophytic life forms. One species, Cymbidium macrorhizon , lacks foliage leaves and has a strongly mycoparasitic existence. Correlation between habitat differentiation and anatomical characters was tested for 21 species of Cymbidium and its putative sister groups. Although hypostomaty characterizes the genus, C. canaliculatum shows amphistomaty. Ecological preference of this species indicates that amphistomaty is likely adapted to intensive insolation. Four types of subepidermal foliar sclerenchyma were found. Two forest floor species, C. goeringii and C. lancifolium as well as the mycoparasitic C. macrorhizon , do not have this sclerenchyma. In this genus, development of sclerenchyma is correlated with the degree of epiphytism. Palisade mesophyll evolved in Cymbidium section Cymbidium . As members of this section grow on isolated trees in tropical lowland forests or on rocks, the differentiation of palisade tissue is probably correlated with immigration to high light habitats. With the exception of C. macrorhizon , stegmata were found in leaves and stems of Cymbidium . Furthermore, a few epiphytic species have stegmata in their roots; this is a curious feature rarely found in vascular plants. Subterranean rhizomes characterize terrestrial species, while ageotropic roots are found in some epiphytic species. Cymbidium macrorhizon shows peculiar features such as degeneration of stomata, anomocytic stomata, and lack of stegmata and sclerenchyma. This set of character transformations is probably correlated with the evolution of mycoparasitic existence. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 138 , 383–419.     

Anatomical structure and distribution of secondary metabolites as a peripheral defence strategy in Aloe hereroensis leaves

Eight leaves from four different plants of Aloe hereroensis and 18 leaf parts of each leaf were tested by anatomic, fluorescence microscopy and TLC methods. Four phenolic secondary metabolites, homonataloin and three isomers of aloeresin, were found in the leaves. The highest content of these metabolites was found in the top third of a leaf and along the leaf margins. In the margins, the content of the four secondary metabolites in the adaxial was higher than in the abaxial direction. In the centre parts of the leaves, the metabolic content of the abaxial parts was higher than in the adaxial parts. The results indicate that homonataloin mainly accumulates in the inner bundle sheath cells (IBSC). The three isomers of aloeresin appear in the outer bundle sheath and in the boundary cells between the chlorenchyma and water-storage tissues. The density of the vascular bundles, the area ratio of the chlorenchyma to the tested counterpart, and the area ratio of the IBSC to a whole bundle are important structural factors to determine the differences in the content of these four secondary metabolites in all the leaf parts. The distribution according to the rosette leaf arrangement and the existence of the 'cocktail' of four phenolic secondary metabolites indicate a peripheral defence strategy of this plant. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 138 , 107–116.     

Alternative modes of leaf dissection in monocotyledons

Although a majority of monocotyledons have simple leaves, pinnately or palmately dissected blades are found in four orders, the Alismatales, Pandanales, Dioscoreales and Arecales. Independent evolutionary origins of leaf dissection are indicated by phylogenetic analyses and are reflected in the diversity of mechanisms employed during leaf development. The mechanism of blastozone fractionation through localized enhancement and suppression of growth of the free margin of the leaf primordium occurs in the Araceae and Dioscoreaceae. By contrast, the corrugated, dissected leaves of palms (Arecaceae) develop through a two-step process: first, plications are formed through intercalary growth in a submarginal position and, second, the initially simple leaf blade is dissected through an abscission-like process of leaflet separation. A third mechanism, perforation formation, is employed in Monstera and five related genera of the Araceae. In this mode, discrete patches of cells undergo programmed cell death during lamina development, resulting in formation of open perforations. When perforations are positioned near the leaf margin, mechanical disruption of the thin bridges of marginal tissue results in a deeply pinnatisect blade. Whereas blastozone fractionation defines the early primary morphogenesis phase of leaf development, the other two modes occur later, during the secondary morphogenesis/histogenesis phase. Evolution of these mechanisms presumably has involved recruitment of other developmental programmes into the development of dissected leaves.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 25–44.     

喀斯特地区莎叶兰的解剖构造及其环境适应性

以广西西北部雅长兰科植物保护区的莎叶兰( Cymbidium cyperifolium)为对象,采用石蜡切片法对莎叶兰叶片和根的解剖构造及其对喀斯特环境的适应性进行了研究。结果表明：(1)莎叶兰叶片的上表皮覆盖有较厚的角质层,气孔均分布于下表皮,且凸出表皮细胞之上;各表皮性状在叶片不同部位存在显著差异,叶片下部的气孔密度、气孔指数和气孔长度最大,表皮细胞密度以叶片上部的最大;叶片属于等面叶,叶肉无栅栏组织和海绵组织的分化;叶脉为明显的平行脉,且粗细交互分布;(2)莎叶兰根的横切面包括根被、皮层、中柱3部分,其中根被细胞排列紧密,为生活细胞;皮层由薄壁细胞组成;根部维管束属于辐射维管束,14原型。菌根粗壮,稀根毛,共生真菌主要分布于根被及皮层中,菌丝体通过根被薄壁细胞间隙及内、外皮层的通道细胞进行侵染。(3)莎叶兰叶片和根的结构不仅有湿生植物特征,如叶片相对较薄、气孔少且凸出表皮细胞、冠/根比值大等;还有旱生植物的特征,如叶片角质层较厚、机械组织发达、细胞结构紧密、具含晶细胞,肉质根具根被,内、外皮层细胞壁明显增厚等。这些结构是莎叶兰对当地缺水、干湿季明显、分布于林下多石砾土壤的生长环境的一种高度适应性表现。     

SEM observations on petals and fruits of some Turkish endemic Bupleurum L. (Umbelliferae) species

Petal and fruit surface sculpturing are reported for seven endemic Bupleurum species in Turkey. Considerable differences in the dorsal surface of petals were observed, but major structural similarities were found in the ventral surface of petals. Significant taxonomic differences between species were determined on the surfaces of the fruits. These microstructural surface features are evaluated as possible consistent parameters in the delimitation of species of Bupleurum. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 138 , 441–449.     

A new species of the Brazilian genus Chresta (Asteraceae, Vernonieae) from Bahia

A new species of the genus Chresta Vell. ex DC. (Vernonieae, Asteraceae) from the Brazilian state of Bahia is described and illustrated. The new species resembles Chresta pinnatifida (Philipson) H.Rob. but differs in having solitary glomerules, leaf bases clasping the stem and 35–40 capitula per glomerule. It can be distinguished from Chresta harleyii and Chresta martii , the other two species of the genus with pinnately lobed leaves, by the type of pubescence, leaf blade bases and florets number. In addition, a key to distinguish all the species of the genus is presented. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 587–590.     

Comparative and functional leaf anatomy of selected Alstroemeriaceae of mainly Chilean origin

The resupinate leaves of 16 species of Alstroemeriaceae were examined using light- and scanning electron microscopy. The leaf anatomy is described for all of the species, that of the petiole and stem for selected species. The mesophyll consists of chlorenchyma and includes idioblasts that contain raphides. Dorsiventral, isolateral or isobilateral leaf types were observed. Petioles are always isolateral. Two epidermal types are commonly observed: type I contains jigsaw puzzle-like intercostal cells and longitudinally elongated cells above the veins; type II contains only longitudinally elongated cells, usually longer, above the veins. Some species have an epidermis which differs from the main types. All species show adaptation to resupination by having an inverted anatomy. Due to the twist of the leaves, adaxial indicates the lower surface and abaxial the upper. Stomata are found on both surfaces. Palisade cells, when they occur, are always on the abaxial surface. Most species are mesomorphic in spite of the dry environments in which they grow; a few show xeromorphic features. The leaves are compared and discussed with relation to ecological conditions.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 261−272.     

Phylloclade anatomy in Phyllanthus section Choretropsis (Phyllanthaceae)

The phylloclade anatomy of nine Brazilian species of the genus Phyllanthus section Choretropsis was studied, and its systematic significance was analysed. The arrangement of epidermal cells varied according to the species, with the guard cells of the stomatal apparatus distributed randomly or with the longitudinal axis parallel to the branch. In both branch types, the stomata were distributed on the phylloclade surface, except in P. gladiatus which showed hypostomatic branches. In most of the species, phylloclades developed a vascular system surrounded by clusters of fibres and cortical parenchyma, which could differentiate into palisade‐like tissue and ground tissue like a true leaf blade. The distribution of the vascular system followed the cylindrical or flattened diameter shape of the branch. In subsection Choretropsis, the cylindrical branches of P. choretroides, contrary to other species, developed a second ring formed by clusters of fibres surrounded by large parenchymal cells. In subsection Applanata, the palisade‐like cells of P. flagelliformis phylloclades acquired an isodiametric shape. The highest level of specialization was observed in P. gladiatus, which developed phylloclades with a dorsiventral structure, a main vein, and small sets of lateral vascular bundles like a typical leaf blade. The tissue arrangement of different species showed a gradual pattern, which was correlated with the different branch morphology and plant architecture. Despite the transitional pattern, it was suggested that some of the phylloclades had an independent origin and convergent evolution in the Choretropsis section. The anatomy of different branch types was shown to be useful, either in isolation or in association with other morphological characters, to delimit the species and to further understand their relationships and phylogeny. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157 , 91–102.     

Presence of Tyloses at the Blade/Sheath Junction in Senescing Leaves of Lolium temulentum L.

Aspects of age-related changes occurring in the blade/sheathjunction of leaves ofLolium temulentum L. were studied usingautoradiography, light and transmission electron microscopy.With increasing leafage the apical transport of sheath-appliedradioactive amino acids was retarded at the sheath side of theblade/sheath junction. An anatomical study revealed the presenceof tyloses in the protoxylcm lacunae of vascular bundles atthis region in senescing leaves. The possible involvement oftyloses in grass leaf senescence is discussed.     

Indeterminate leaves of Chisocheton (Meliaceae): survey of structure and development

Pinnately compound leaves in the Malesian genus Chisocheton (Meliaceae) have leaf-tip buds that continue to produce new pinnae (leaflets) periodically for many years. Juvenile leaves form a terminal pinna in place of the leaf-tip bud found in adult leaves. The histology of an old leaf-tip bud is similar to the entire leaf primordium in other species of Meliaceae with large pinnate leaves (e.g. Chukrasia and Dysoxylum ) which serve as examples of more typical leaves. Pinna initiation from this meristem continues after the first stage of leaf expansion as seen in the relatively constant number of pinna primordia in a large sampling of leaf-tip buds of varying ages. Structure and development are compared in leaves of nine species of Chisocheton , out of a total of approxiamtely 50 species in the genus. Species having small leaves (e.g. C. pentandrus ) show more branch-like, indeterminate leaf growth as compared with species with large leaves (e.g. C. macranthus ). The structure and development of leaves of Chisocheton are like the similar indeterminate leaves of the American and African genus Guarea . Some authors have used the indeterminate leaves of Chisocheton and Guarea as examples of intermediate organs showing 'fuzzy morphology' or 'partial homology.' Nevertheless, these unusual organs are considered here as being homologous with leaves of other Meliaceae based on their position, histology and ontogeny. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 207–221.     

Ultrastructural studies on the process of aloin production and accumulation in Aloe arborescens (Asphodelaceae) leaves

Aloin, a kind of anthraquinone, is a chemical component in Aloe leaves used in medicine. The processes of aloin production, transport and storage were studied with a transmission electron microscope using the lead acetate precipitate method for ultracytochemical localization of aloin in the leaf of Aloe arborescens Mill. Results showed that aloin was produced in the plastids of the assimilating tissue, transported through the plastid membrane to the surrounding endoplasmic reticulum and enveloped in the vesicles by the endoplasmic reticulum elements. The vesicles approached, and later fused with, the plasmalemma, released their contents into the apoplast through exocytosis and finally, reached the vascular bundle sheath by apoplastic translocation. Aloin was transported to the internal tangential wall of the vascular bundle sheath cell through endoplasmic reticulum vesicles, and reached the cytoplasm of the aloin cell by means of plasmodesmata. Finally, aloin was stored in the vacuole of the cell in which it was produced.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 241–247.