Exogenous Methyl Jasmonate Alters Trichome Density on Leaf Surfaces of Rhodes Grass (Chloris gayana Kunth)

Jasmonates, including jasmonic acid and its derivatives such as methyl jasmonate (MeJA), are plant growth substances that control various responses. Jasmonates regulate leaf trichome density in dicotyledonous plants, but their effects on the trichome density of monocotyledonous plants, such as those in the Poaceae, remain unclear. In the present study we examined the effects of exogenous MeJA on the trichome density of Rhodes grass, which has three kinds of trichomes: macrohairs, salt glands, and prickles. Exogenous MeJA significantly increased the densities of macrohairs and salt glands on the adaxial and abaxial leaf surfaces and those of prickles on the adaxial leaf surface. Because exogenous MeJA significantly reduced the leaf area, we calculated the number of trichomes per 1000 epidermal cells to eliminate the effects of reduced leaf area. Exogenous MeJA significantly increased the number of macrohairs per 1000 epidermal cells on both adaxial and abaxial leaf surfaces, but it significantly decreased the number of salt glands per 1000 epidermal cells on both surfaces. Exogenous MeJA had no significant effects on the number of prickles per 1000 epidermal cells on either of the leaf surfaces. These results indicate that exogenous MeJA alters the trichome density by affecting leaf area and trichome initiation, and the effects of exogenous MeJA on trichome initiation differ among the various trichome types.     

禾本科广义拂子茅属的叶表皮形态研究

禾本科广义拂子茅属Calamagrostis Adans. s.l.是广布于全球温带和热带亚热带高山的一个大属, 常分为拂子茅属Calamagrostis Adans. s.s.和野青茅属Deyeuxia Beauv.。对国产5种拂子茅属和26种、1变种野青茅属植物在光镜下进行了叶表皮特征的观察。发现广义拂子茅属植物的叶表皮特征为典型的狐茅型, 很多结构在种间有丰富的变异式样, 如脉间长细胞形状、大小和细胞壁的厚度与弯曲程度, 短细胞形状和分布式样, 硅质体形状和分布式样, 气孔形状和分布式样, 以及刺毛形态和分布式样等。在国产种类中首次发现微乳突结构, 在很多种类中发现刺毛硅质化现象。用UPGMA对15个叶表皮性状进行分析, 结果分为两大支: 具加厚的长细胞、密集分布的短细胞和(或)刺毛以及刺毛硅质化的种类聚为一支; 具薄壁的长细胞、较稀疏的短细胞和(或)刺毛以及刺毛不发生硅质化的种类聚为另一支。这种分异与广义拂子茅属的属下系统相关性不大, 但与种的海拔分布有关。前一支的种类大多生长于高海拔(2600 m以上)地区, 而后一支的种类则大多生长在低海拔(2600 m以下)地区。     

中国看麦娘属植物叶表皮微形态研究

看麦娘属是禾本科早熟禾亚科的重要类群,中国分布有8种。该研究采用光学显微镜和电子扫描电镜,对中国看麦娘属7种植物的叶下表皮(含叶鞘下表皮)微形态特征进行观察并探讨其分类学意义。结果表明：(1)看麦娘属叶鞘下表皮长细胞的形状呈长筒状或短筒状,深波状弯曲的细胞壁,脉间短细胞密布,单生或对生,形状多为新月形或椭圆形,脉区硅细胞为椭圆形,平行形或低圆屋顶形的气孔器,看麦娘属叶鞘下表皮特征高度一致,分类价值不大。(2)看麦娘属植物叶下表皮和叶鞘下表皮的显微结构存在很大不同,看麦娘属植物叶下表皮长细胞多为长筒状或短筒状或长菱形,平周壁平直或不明显弯曲,脉间区短细胞分布或无,脉区刺毛密布,脉间区有或无刺细胞的分布,平行形的气孔器,脉区分布的硅细胞多为长方形边缘呈波状弯曲。看麦娘属植物叶下表皮的不同主要体现在脉间区长细胞的形状及平周壁的弯曲与否,脉间区短细胞出现或无,脉间区刺细胞的分布数量以及气孔器的列数。7种植物的叶表皮微形态的分类检索表据此列出。依据叶表皮微形态划分的组与根据外部特征划分的组并不完全一致。但是,由于该文仅研究了7种,叶表皮微形态对于看麦娘属的分类意义还需要更深入的研究。     

Micromorphology of the leaf epidermis in taxa of the Agropyron-Elymus complex (Poaceae)

WEBB, M. E. & ALMEIDA, M. T., 1990. Micromorphology of the leaf epidermis in taxa of the Agropyron-Elymus complex (Poaceae). A comparative analysis by scanning electron microscopy was carried out on both leaf epidermes (adaxial and abaxial) of Elymus pychnanthus (Godr.) Meld, and Agropyron glaucum Roemer & Schultes.
The adaxial epidermes of E.pychnanthus and A. glaucum are similar in the position and the shape of the long cells, silica bodies and costal and papillate prickles. They differ in the higher number of silica cells in E. pychnanthus and in the presence of intercostal hooks in A. glaucum.
The abaxial epidermes of both species are similar in the shape of interstomatal cells, silica bodies and papillate prickles. They differ in the distribution of the stomata and in the contact zones of the long cells. Elymus pychnanthus and A. glaucum differ also in the trichomes situated along the margins of the leaf blade.
These micromorphological differences, especially those of the abaxial epidermis, are useful taxonomic features.     

Effects of plant spinescence on large mammalian herbivores

Summary Plant thorns and spines had these effects on the feeding behaviour of the three species of browsing ungulate that we studied, kudu, impala and domestic goats: (i) bite sizes were restricted, in most cases to single leaves or leaf clusters; (ii) hooked thorns retarded biting rates; (iii) the acceptability of those plant species offering small leaf size in conjunction with prickles was lower, at least for the kudus, than those of other palatable plant species; (iv) the inhibitory effect of prickles on feeding was much less for the smaller impalas and goats than for the larger kudus; (v) from certain hook-thorned species the kudus bit off shoot ends despite their prickles; (vi) for certain straight-thorned species the kudus compensated partially for the slow eating rates obtained by extending their feeding durations per encounter. Most spinescent species were similar in their acceptability to the ungulates to unarmed palatable species, despite higher crude protein contents in their foliage than the latter. Such structural features furthermore reduce the tissue losses incurred by plants per encounter by a large ungulate herbivore, by restricting the eating rates that the animals obtain. In this way prickles function to restrict foliage losses to large herbivores below the levels that might otherwise occur.     

中国杜鹃花属的叶表皮特征及其系统学意义

通过光学显微镜和扫描电镜对国产杜鹃花属Rhododendron6个亚属48种4变种植物的叶表皮特征进行观察,结果表明叶表皮的鳞片、腺体、表皮毛的有无和气孔器的类型具有一定的系统学意义。光学显微镜下可见：叶表皮细胞形状为多边形、不规则形,垂周壁式样为较平直、浅波状和波状．气孔器普遍存在于下表皮,上表皮未见气孔器分布,其类型均为无规则形。杜鹃亚属subgen．Rhododendron叶表皮具有鳞片,在扫描电镜下可见气孔器散生于指状突起之间,有别于无鳞类杜鹃花。常绿杜鹃亚属subgen．Hymenanthes和映山红亚属subgen．Tsutsusi有多种不同叶表皮类型;除了凯里杜鹃R．westlandii和弯蒴杜鹃R．henryi形态较特殊之外,马银花亚属subgen．Azaleastrum的马银花组sect．Azaleastrum和长蕊杜鹃组sect．Choniastrum微形态区别不明显。羊踯躅亚属subgen．Pentanthera的羊踯躅R．molle叶表皮有表皮毛,无腺体,气孔器密生,外拱盖形状不对称,外围有多层波形条纹突起可与其他亚属区分：而叶状苞亚属subgen．Therorhodion的叶状苞杜鹃R．redowskianum保卫细胞有明显的T型加厚,与其他亚属种类明显不同,与分子系统发育研究得到的“叶状苞亚属是杜鹃花属的基部类群”的结果吻合。本研究还依据叶表皮微形态特征讨论了一些近缘种类的关系。     

Leaf anatomy as a contribution to the taxonomy of Salacioideae N.Hallé ex Thorne & Reveal (Celastraceae)

The current classification systems recognize Salacioideae as a monophyletic group within Celastraceae. Nonetheless, some divergences exist for genera: in some cases, most species of the subfamily have been included in only two genera; in others, these genera have been subdivided. This study characterizes the leaf anatomy of 31 species of the subfamily Salacioideae as a contribution to identifying them through features that may also help distinguish among genera. Cross-sections of the median region of the leaf blade and of the petiole and dissociated and macerated epidermis were analyzed. Taxonomically relevant anatomical characters include the type of crystals in the parenchymatous tissue (monocrystals in Cheiloclinium and druses in other genera); the presence of laticifers in Cheiloclinium and Tontelea only; the variable form of the petiole vascular system among studied species; the type of stomata (cyclocytic with two concentric circles of subsidiary cells in P. dulcis; anomocytic in T. attenuata, T. fluminensis, and T. leptophylla; laterocytic in C. anomalum and C. hippocrateoides; and ciclocytic in the other species); the sinuosity of the anticlinal walls of the epidermal cells (sinuous in Cheiloclinium and Peritassa, except P. laevigata, and in S. arborea, S. insignis, S. mosenii, S. nemerosa, and S. opacifolia, and straight in all other studied species); the presence of crystalliferous idioblasts in the epidermis of P. dulcis, P. flaviflora, and P. mexiae; and the presence, form, and disposition of sclereids in the leaf blade, which is a highly variable character among the studied species.     

中国独行菜族部分属种的叶表皮观察

利用光学显微镜对中国十字花科独行菜族部分广布属及相关属植物进行了叶表皮微形态学观察研究,结果表明:所研究的中国独行菜族植物叶表皮微形态具有多样性,可以划分为两种类型:第一类,叶上下表皮同形或近同形,叶表面被单毛或光滑无毛,包括独行菜属、群心菜属、菘蓝属和菥蓂属;第二类,叶上下表皮不同形,叶表面被分支毛和单毛,包括荠属和亚麻荠属;叶表皮微形态结果支持荠属应从独行菜族中移出,独行菜族是一多系类群,同时认为荠属与亚麻荠属植物具有较近的亲缘关系。     

Leaf epidermal features of Rhododendron (Ericaceae)from China and their systematic significance

Rhododendron is the largest genus within the subfamily Rhododendroideae, which has about 1000 known species in the world and more than 500 species in China. Since the genus was established by Linnaeus, its infrageneric relationships have been well studied by many taxonomists on the basis of morphological characters and molecular data. In 1996, Chamberlain et al. proposed a new system of Rhododendron with eight subgenera, i.e., Azaleastrum, Candidastrum, Hymenanthes, Mumeazalea, Pentanthera, Rhododendron, Therorhodion, and Tsutsusi. In this paper, micromorphological characters of leaf epidermis in 4 varieties, 48 species, 6 subgenera of Rhododendron from China were examined using light microscopy (LM) and scanning electron microscopy (SEM). Leaf epidermal features are described and micromorphological types are distinguished here according to morphological characters such as scale, gland, foliar trichome and stomatal apparatus of leaf epidermis. It is shown that the leaf epidermal cells are usually irregular or polygonal in shape. The patterns of anticlinal walls are straight, arched or undulate. The stomatal apparatuses are anomocytic and are usually found on abaxial, not adaxial, epidermis. The results also show that: (1) the lepidote rhododendron (i.e., subgen. Rhododendron), which has both scales and papillae on leaf epidermis, differs distinctly from the elepidote rhododendron; (2) three types of leaf epidermis are identified in subgen. Hymenanthes (i.e., R. fortunei-type, R. chihsinianum-type and R. simiarum-type), whereas four in subgen. Tsutsusi (i.e., R. mariesii-type, R. simsii-type, R. mariae-type and R. flosculum-type); (3) except for R. westlandii and R. henryi, the species of subgen. Azaleastrum show similar morphological characters, i.e., dense stomatal apparatuses surrounded by ringed or discontinuous striates; (4) R. molle of subgen. Pentanthera differs from the species of other subgenera on morphological characters such as foliar trichomes, dense stomatal apparatuses with asymmetrical outer stomatal rims surrounded by undulate-striates, and no gland; (5) only R. redowskianum is found with distinct T-pieces at the polar region of guard cells in Rhododendron. The results support the conclusion inferred from molecular systematic studies that subgen. Therorhodion is the basal clade of Rhododendron. Finally, the relationships between the closely related species are also discussed on the basis of leaf epidermal features.     

中国29种和泰国1种卷柏科植物的叶形态学研究及其分类学意义

利用光学显微镜和体视显微镜对卷柏科Selaginellaceae中国产29种和泰国产1种植物的叶形态进行了宏观和微观观察。结果表明卷柏科植物在叶形态学方面表现出较高的一致性。(1)叶多为二形, 罕为一形, 无叶柄, 叶二形者侧叶多大于中叶, 外展或斜上, 中叶贴生于腹面茎上, 交互覆瓦状排成4列, 叶一形者则在茎上作螺旋状排列。(2)侧叶与中叶上表皮细胞形态相似, 近等径方形或长方形, 垂周壁平直、弓形或波状; 侧叶与中叶下表皮细胞形态也相似, 长方形或长条形, 垂周壁波状。(3)气孔类型为无规则型或放射型, 气孔多分布于中叶上表皮和侧叶下表皮中脉处。研究发现, 叶外形、叶表皮细胞形态及气孔分布特征等为中国卷柏科植物种间分类提供了重要的参考依据。 研究的有关种类可以依据下面这些特征进行分类: (1)叶为一形叶或二形; (2)叶在枝上着生为螺旋状或成对交互排列; (3)叶表皮细胞垂周壁为平直、弓形、深波状或浅波状; (4)叶上、下表皮细胞同形或异形; (5)气孔为无规则型或放射型; 以及(6)气孔在中叶和侧叶的上、下表皮上的分布等特征。     

麦冬、土麦冬和阔叶土麦冬叶表皮形态结构的观察

用光镜和扫描电镜观察了麦冬（Ophiopogon japonicus（L．f）Ker—Gawl.]、土麦冬（Liriope spicata Lour．）和阔叶土麦冬（L．platyphylla Wanget Tang）叶表皮显微结构、亚显微结构和角质层内表面的形态结构。结果表明,气孔主要分布于麦冬、土麦冬和阔叶土麦冬叶片的下表皮,气孔密度分别为76．4、114．3和99．8个·mm^-2;仅阔叶土麦冬叶片上表皮有少量气孔分布。3种植物的气孔器均不具有副卫细胞,并在叶脉间形成纵向气孔带。表皮细胞长方形,气孔带与非气孔带处表皮细胞的形态和大小差异较明显。麦冬气孔周围的表皮细胞平周壁具明显瘤状突起,导致气孔下陷;土麦冬气孔周围的表皮细胞平周壁呈波浪状突起,使气孔相对下陷;阔叶土麦冬气孔周围的表皮细胞平周壁基本无突起,气孔不下陷。3种植物的叶表皮均有发达的角质层和丰富的蜡质,且蜡质主要分布于下表皮气孔带处。这些结构特征可能与它们所具有的喜阳、耐阴和耐旱等特性有一定的相关性。     

Silica bodies and hooked papillae in lemmas of Melica species (Gramineae: Pooideae)

Melica (sensu lato ) is a genus with approximately 80 species distributed in the temperate regions of both hemispheres. The genus has been better studied morphologically than anatomically, but morphological patterns have led to conflicting subdivisions either into sections, or by segregation of the genus Bromelica . The lemma epidermis is examined here under scanning electron microscopy (SEM) for 40 species of Melica to see if it exhibits characters of systematic significance. Silica bodies and hooked papillae are reported in the lemma for the first time in the genus. Patterns of variation are evident within four structures that terminate in points: prickles, hooks, hooked papillae and macro-hairs. Fourteen qualitative characters for lemma epidermis are used in phenetic analyses. The presence or absence of macro-hairs conflicts with some other patterns, but gives some support to the classification of Papp. Characters associated with the split of the clusters were micro-hairs (straight and geniculate) and the distribution and frequency of prickles in both costal and intercostal zones. As a conclusion, Melica is not a coherent taxonomic grouping and a simple segregation of Bromelica from Melica genus is rejected.  © 2003 The Linnean Society of London , Botanical Journal of the Linnean Society , 2003, 141, 447 – 463.     

中国特有属藤山柳属(猕猴桃科)植物的叶表皮形态及其分类学意义

由于形态特征变异和地理分布区域存在重叠,中国特有属藤山柳属(猕猴桃科)的物种划分问题长期以来存在争议,曾被分为20种或修订为含1种4个亚种的单型属。该研究选取了在形态和地理分布上有代表性的29个居群的184份标本,利用光学显微镜和扫描电镜观察了叶表皮形态和微形态特征,以探讨它们的分类学意义。结果表明:(1)藤山柳属植物叶表皮毛被的形态和微形态特征有3类,即光滑-短柱状毛、刚毛-长柱状毛/长刺毛、绒毛-单列多细胞毛,且这些特征在居群间差异明显,并各具明显的地理分布格局,支持把藤山柳属分为3类,即光滑类、刚毛类和绒毛类。(2)3类藤山柳植物在个别居群表现出部分同域分布现象,在峨眉山不同海拔高度的3个居群存在垂直地带性分布特点。(3)藤山柳属植物叶表皮的其他形态特征,如不规则型表皮细胞、6类气孔器、叶表皮初级蜡质纹饰以网状隆起为主,伴随着2～4类次级纹饰,在居群间变化多样等均没有明显的分类学意义。(4)由于具有相同的叶表皮形态特征和地理分布,建议把繁花藤山柳合并到绒毛藤山柳,故支持藤山柳属是1个正处于分化进程中的单型属,包括1个种3个亚种。     

Rose Prickles and Asparagus Spines – Different Hook Structures as Attachment Devices in Climbing Plants

Functional morphology and biomechanical properties of hook structures functioning as attachment devices in the leaning climbers Rosa arvensis, Rosa arvensis ‘Splendens‘, Asparagus falcatus and Asparagus setaceus are analysed in order to investigate the variability in closely related species as well as convergent developments of hook structure and properties in distant systematic lineages (monocots and dicots). Prickles and spines were characterised by their size, orientation and the maximum force measured at failure in mechanical tests performed with traction forces applied at different angles. In Rosa arvensis and Rosa arvensis ‘Splendens‘ three types of prickles differing largely in geometrical and mechanical properties are identified (prickles of the wild species and two types of prickles in the cultivar). In prickles of Rosa arvensis no particular orientation of the prickle tip is found whereas in the cultivar Rosa arvensis ‘Splendens‘ prickles gradually gain a downward-orientation due to differential growth in the first weeks of their development. Differences in mechanical properties and modes of failure are correlated to geometrical parameters. In Asparagus falcatus and Asparagus setaceus spines are composed of leaf tissue, stem tissue and tissue of the axillary bud. Between species spines differ in size, orientation, distribution along the stem, tissue contributions and mechanical properties. The prickles of Rosa arvensis and its cultivar and the spines of the studied Asparagus species have several traits in common: (1) a gradual change of cell size and cell wall thickness, with larger cells in the centre and smaller thick-walled cells at the periphery of the hooks, (2) occurrence of a diversity of shape and geometry within one individual, (3) failure of single hooks when submitted to moderate mechanical stresses (F_max/basal area < 35 N/mm²) and (4) failure of the hooks without severe stem damage (at least in the tested wild species).     

A Developmental Study of Silicification in the Trichomes and Associated Epidermal Structures of the Inflorescence Bracts of the Grass, Phalaris canariensis L

Silicon accumulation is greatest in the abaxial epidermis ofthe inflorescence bracts of Phalaris canariensis L. It occursinitially in glumes 1 week before emergence, at low levels inthe trichomes and long cells, but at high levels in the papillae.Accumulation in long cells remains low throughout, althoughit increases with time. Papillae and prickle hair silicon countsconsiderably exceed values for other epidermal structures. Siliconis also deposited in glume macrohairs before emergence, andat maturity reaches intermediate levels. Little silicon is present in the macrohairs and long cells ofthe abaxial epidermis of the fertile lemma prior to emergence.After emergence, however, silicon is rapidly accumulated inthese structures, and at maturity they have the highest siliconlevels of all the mature bract tissues. Silicon accumulation in the lemma lags behind that of the glumeuntil emergence, when it increases more rapidly, and by maturityexceeds that of the glume in macrohairs and long cells. Thesignificance of the results is discussed in relation to wallthickening, bract anatomy and panicle emergence. Phalaris canariensis, L, canary grass, inflorescence bracts, silicification, trichome, glume, lemma     

Grasses of different C4 subtypes reveal leaf traits related to drought tolerance in their natural habitats: Changes in structure, water potential, and amino acid content

Three grasses (Poaceae) of different C(4) subtypes, Paspalum dilatatum (NADP-malic enzyme [ME]), Cynodon dactylon (NAD-ME) and Zoysia japonica (phosphoenolpyruvate carboxykinase), occur in natural habitats that differ in annual rainfall. Their leaf characteristics were studied to identify traits related to drought tolerance. Plants were grown in pots, and water deficit was gradually induced by withholding water. Leaves of Z. japonica had the greatest and P. dilatatum the lowest relative dry matter content. Transverse sections of leaves that developed during the water deficit showed little change compared to control leaves, consistent with low phenotypic plasticity. Anatomical features distinguished the three species, with xeromorphic characteristics most strongly represented in Z. japonica. The leaf relative water content (RWC) decreased with the soil water content similarly for the three grasses. However, at 80% RWC, the leaf water potential was -3.1 MPa for Z. japonica and only -1.3 MPa for P. dilatatum and C. dactylon. Soluble amino acids, especially proline, increased as RWC decreased in leaves of C. dactylon and Z. japonica. Phenylalanine, valine, leucine, and isoleucine increased more in Z. japonica than in the other two species. The results provide evidence that C. dactylon and, especially, Z. japonica have evolved leaf traits better suited to arid habitats.     

Leaf epidermal morphology of Zanthoxylum s.l. (Rutaceae) from China

Leaf epidermis characters in 40 of ca. 250 species of Zanthoxylum (Rutaceae) were investigated using light microscopy (LM) and scanning electron microscopy (SEM). The stomata are anomocytic and exist only on the abaxial epidermis except in Z. nitidum, which also has stomata on the adaxial surface. The epidermal cells are usually polygonal or irregular in shape, with anticlinal walls straight, arched, repand, or sinuous. Under the SEM, the inner margin of the outer stomatal rim is nearly smooth, sinuolate or erose, and the cuticular membrane of the leaf epidermis is smooth, striate, or sometimes striate to wrinkled. These data of leaf epidermis of Zanthoxylum demonstrated that there exist many common characters between subgen. Fagara and subgen. Zanthoxylum, sug-gesting a close relationship between the two subgenera. The utility of some characters in identifying some species of Zanthoxylum was also discussed.     

The maize macrohairless1 locus specifically promotes leaf blade macrohair initiation and responds to factors regulating leaf identity

The leaf surfaces of almost all plant species possess specialized epidermal cell types that form hairs or trichomes. Maize leaves produce three distinct types of hairs, the most prominent being the macrohairs that serve as a marker for adult leaf identity and may contribute to insect resistance. This report describes the maize macrohairless1 (mhl1) locus, which promotes macrohair initiation specifically in the leaf blade. Each of seven recessive mhl1 mutant alleles significantly reduces or eliminates macrohairs in the leaf blade. The mhl1 mutations block macrohair initiation rather than interfering with macrohair morphogenesis. Genetic mapping placed mhl1 within bin 4 on chromosome 9. A second independently segregating locus was found to partially suppress the mhl1 mutant phenotype in certain genetic backgrounds. Macrohair density was observed to increase during early adult vegetative development and then progressively decline, suggesting macrohair initiation frequency is affected by factors that act throughout shoot development. Genetic analyses demonstrated that mhl1 acts in the same pathway but downstream of factors that either promote or repress adult leaf identity. Thus, mhl1 plays a key role in integrating developmental programs that regulate leaf identity during shoot development with those that specify macrohair initiation within the leaf blade.