A comparative study on the anatomy and development of different shapes of domatia in Cinnamomum camphora (Lauraceae)

BACKGROUND AND AIMS: Domatia are small organs usually found in the axils of major veins on the underside of leaves and, although they have received wide attention from ecologists, few detailed reports exist on their anatomy or development. This study is focused on the domatia of Cinnamomum camphora (Lauraceae) and is the first comparative study on the anatomy and development of the different shapes of domatia within a single plant. METHODS: Four types of domatia in C. camphora leaves were observed on paraffin sections under a microscope. KEY RESULTS: The domatia consisted of six histological parts: the upper epidermis, the upper mesophyll tissue, spongy tissue, the lower mesophyll tissue, the tissue filling the rim opening, and the lower epidermis. They differed from the non-domatial lamina mainly in the cell structure of the upper and lower mesophyll tissue and the rim tissue. Differences in domatium shapes were mainly associated with differences in the structure of the upper mesophyll and in the number and size of the rim tissue cells. Differences in the development of domatium types were observed in terms of initiation timing, differentiation of the upper mesophyll cells and degree of rim tissue development. CONCLUSIONS: In domatia, active anticlinal division in the lower mesophyll cells, as compared with the upper mesophyll cells, was coordinated with dynamic growth of rim tissue cells and resulted in cavity formation. The anatomical or developmental differences among the four types of domatia were related to the positions of the domatia within a leaf. In terms of the ecological implications, the major anatomical difference between the domatia used by herbivorous and carnivorous mites was in the development of the rim tissue.     

OCCURRENCE OF PREDATORY AND FUNGIVOROUS MITES IN LEAF DOMATIA

Domatia are small invaginations and hair tufts usually found at vein junctions on the undersides of leaves in many woody dicots. Domatia of 32 plant species (of worldwide origin from 18 families) growing in California, Hawaii, and Costa Rica were examined for mites. Domatia of 31 of 32 (97%) of these plants contained mites, and 24 of 32 (75%) contained mite eggs. Mites were found within the domatia of 48% of the sampled leaves. The domatia of 26 of 31 (84%) plants had mite species considered beneficial (primarily in the families Phytoseiidae and Tydeidae, but also Bdellidae, Cheyletidae and Stigmaeidae), while 6 of 31 (19%) had mite species considered harmful (Tenuipalpidae and Eriophyidae). Based on these findings (and in part upon Lundström's 1887 domatia theory), we hypothesize the existence of a widespread facultative mutualism between plants with leaf domatia and beneficial mites: leaf domatia serve as shelters and nurseries for mites which in turn eat phytophagous arthropods and pathogens using the plants. This proposed mutualism could be of importance to agriculture since domatia are known to occur in some crop plants, including coffee, grape, and walnut.     

Leaf domatia in Korean plants: floristics,frequency, and biogeography

Leaf domatia, specialized structures on the undersides of leaves, appear to promote mutualism between plants and those mites that protect leaves from pathogenic fungi and small herbivorous arthropods. Their occurrence was surveyed on 425 plant species (368 native species and 57 species introduced from other regions of northeastern Asia), comprising 177 genera and 59 families of woody Dicotyledonae in the Korean flora. Domatia, primarily of tuft or pocket forms (84% of species), were present in vein axils on leaves of 152 species (36%) within 66 genera (37%) and 39 families (66%). Overall, we report leaf domatia in nine new families and 25 additional genera. Mites, primarily from families or suborders (e.g., Acaridida, Phytoseiidae, Tydeidae) in which arboreal representatives are primarily fungivorous or predaceous, were found in domatia on leaves of 78% of all species sampled. Domatia frequency depended upon both growth form and habit of species. They were especially common among tree species (55%), but less so on shrubs (20%) and woody vines (22%). Domatia were much more frequent on species of deciduous plants (40% of those sampled) than on evergreen species, of which only 10% had leaf domatia. This difference was consistent across a number of local deciduous and broadleaf evergreen forests in South Korea. The high frequency of leaf domatia in this survey indicates that plant-mite mutualism may be widespread in Korean plant communities. The floristic affinities of plants with domatia in Korea to those in other regions of East Asia, eastern North America, and Europe suggest that these plant-mite associations characterize the temperate broadleaf deciduous forest biome in the Northern Hemisphere.     

Domatia reduce larval cannibalism in predatory mites

Abstract.  1. Acarodomatia are small structures on the underside of leaves of many plant species, which are mainly inhabited by carnivorous and fungivorous mites.
2. Domatia are thought to protect these mites against adverse environmental conditions and against predation. They are considered as an indirect plant defence; they provide shelter to predators and fungivores and these in turn protect the plants against herbivores and fungi.
3. We studied the possible role of domatia of coffee ( Coffea arabica L.) (Rubiaceae) and sweet pepper ( Capsicum annum L.) (Solanaceae) in reducing cannibalism in the mites inhabiting the domatia. We measured cannibalism of larvae by adults of the predatory mites Iphiseiodes zuluagai Denmark & Muma and Amblyseius herbicolus Chant on coffee leaf discs and of the predatory mite Iphiseius degenerans (Berl.) on sweet pepper leaf. Domatia were closed with glue or left open.
4. Cannibalism in all three species increased when domatia were closed. With I. degenerans , moreover, we found that the previous diet of the cannibal attenuated the effect of domatia on cannibalism.
5. We conclude that domatia can protect young predatory mites against cannibalism by adults and that the diet of cannibals affects the rate of cannibalism.     

Leaf domatia and foliar mite abundance in broadleaf deciduous forest of north Asia

Plant morphology may be shaped, in part, by the third trophic level. Leaf domatia, minute enclosures usually in vein axils on the leaf underside, may provide the basis for protective mutualism between plants and mites. Domatia are particularly frequent among species of trees, shrubs, and vines in the temperate broadleaf deciduous forests in north Asia where they may be important in determining the distribution and abundance of mites in the forest canopy. In lowland and montane broadleaf deciduous forests at Kwangn;akung and Chumbongsan in Korea, we found that approximately half of all woody species in all forest strata, including many dominant trees, have leaf domatia. Pooling across 24 plant species at the two sites, mites occupied a mode of 60% (range 20-100%) of domatia and used them for shelter, egg-laying, and development. On average, 70% of all active mites and 85% of mite eggs on leaves were found in domatia; over three-quarters of these were potentially beneficial to their hosts. Further, mite abundance and reproduction (expressed as the proportion of mites at the egg stage) were significantly greater on leaves of species with domatia than those without domatia in both forests. Effects of domatia on mite abundance were significant only for predaceous and fungivorous mite taxa; herbivore numbers did not differ significantly between leaves of species with and without domatia. Comparable patterns in broadleaf deciduous forest in North America and other biogeographic regions suggest that the effect of leaf domatia on foliar mite abundance is general. These results are consistent with several predictions of mutualism between plants and mites, and indicate that protective mutualisms may be frequent in the temperate zone.     

Cuticular anatomy of Sphenobaiera huangii (Ginkgoales) from the Lower Jurassic of Hubei, China

Sphenobaiera huangii (Sze) Hsü is typical Early Mesozoic fossil foliage of Ginkgoales in China. It has been recorded from the Upper Triassic to the Lower Jurassic. The cuticular anatomy is investigated based on material from the type locality, Lower Jurassic Hsiangchi Formation, Zigui County, Hubei Province. The specimens are similar to S. huangii, but contain new information about leaf morphology and cuticular anatomy. Lower and upper cuticle is investigated using light and electron microscopy (LM, SEM, and TEM). Many features are described for the first time, including general structures of lower and upper cuticle, stomata, papillae, and cuticular ultrastructure. At the ultrastructural level, two layers have been distinguished in both lower and upper cuticle, including a homogeneous outer layer with granules and a heterogeneous inner layer with fibrils. Based on a literature comparison between S. huangii and other relevant species of Sphenobaiera, S. huangii may represent the best-known taxon in the genus Sphenobaiera in both leaf morphology and cuticular structures. This study provides the first detailed ultrastructural data on the leaf cuticle of Sphenobaiera, one of the oldest foliage taxa of Ginkgoales, and offers further evidence for potential discussion on the taxonomic relationships of S. huangii with other ginkgoalean taxa.     

Plants,mites and mutualism: leaf domatia and the abundance and reproduction of mites on Viburnum tinus (Caprifoliaceae)

Associations between mites and leaf domatia have been widely reported, but little is known about their consequences for either plants or mites. By excising domatia from leaves of the laureltinus, Viburnum tinus L. (Caprifoliaceae), in the garden and laboratory, we showed that domatia alter the abundance, distribution, and reproduction of potential plant mutualists. Over 4 months, leaves with domatia on six garden shrubs had 2–36 times more predatory and microbivorous mites, and more mite eggs than leaves without domatia. However, this effect varied among plants and was weaker on one shrub with few mites on its leaves. Domatia also influenced the distribution of mites on leaves. A significantly higher fraction of mites, representing all life stages, was found in vein axils of leaves with domatia than in vein axils on leaves without domatia. Single-leaf experiments in the laboratory showed that domatia enhanced reproduction by the predatory mite, Metaseiulus occidentalis, especially at low relative humidity (30–38%). When domatia were removed, oviposition was reduced significantly only at low relative humidity, suggesting that domatia provide mites with refuge from environmental extremes on the leaf surface. Moreover, the use of domatia by predatory mites may reduce the impact of some plant enemies. In two experiments where prey consumption was measured, M. occidentalis ate significantly higher percentages of the eggs of the two-spotted spider mite (Tetranychus urticae). Our results are consistent with the viewpoint that mite-domatia associations are mutualistic. By directly aiding and abetting the third trophic level, plants with leaf domatia may increase the efficiency of some predaceous and microbivorous mites in consuming plant enemies.     

Ants Induce Domatia in a Rain Forest Tree (Vochysia vismiaefolia)1

In Amazonian rain forest trees of Vochysia vismiaefolia (Vochysiaceae), ants were found to induce twig structures that resembled classical ant domatia. This phenomenon is novel for ant‐plants, which commonly develop domatia without the activity of ants. Eight species of ants were recorded inside the domatia of six individual trees, but domatia were most numerous and characteristic when induced and inhabited by an undescribed species of Pseudomyrmex on two trees. To investigate the mechanism of domatium growth, we drilled holes into young twigs: the expansion of the twig diameter surrounding the holes was significantly accelerated, comparable to domatia formation. Domatia induction is discussed as a putative step in the evolution of ant‐plants.     

安徽产大戟属植物叶表皮微形态

利用光镜和扫描电镜对安徽7种大戟属(Euphorbia)植物叶表皮进行了观察。发现该属7种植物叶表皮细胞为不规则形或多边形,垂周壁一般为平直、浅波状、深波状;气孔器通常为无规则型,有的种类叶表皮具单细胞毛或多细胞毛,表皮毛在电镜下具瘤状突起的纹饰。在扫描电镜下,角质层纹饰多具鳞片或颗粒等特征,种间有细微差别。结果表明该7种植物叶表皮微形态有明显差别,这些表皮特征有助于识别一些在外部形态较相近的种类。     

Structural characters of leaf epidermis and their systematic significance in Vitaceae

Leaf epidermis of 37species representing 11 genera of Vitaceae was investigated using light microscopy（LM）and scanning electron microscopy（SEM）.The shapes of leaf epidermal cells are usually irregular or polygonal;the patterns of anticlinal walls are stra     

Phylogenetic and Experimental Tests of Interactions among Mutualistic Plant Defense Traits in Viburnum (Adoxaceae)

Abstract Plant traits that mediate mutualistic interactions are widespread, yet few studies have linked their macroevolutionary patterns with the ecological interactions they mediate. Here we merged phylogenetic and experimental approaches to investigate the evolution of two common mutualistic plant traits, extrafloral nectaries (EFNs) and leaf domatia. By using the flowering plant clade Viburnum, we tested whether macroevolutionary patterns support adaptive hypotheses and conducted field surveys and manipulative experiments to examine whether ecological interactions are concordant with evolutionary predictions. Phylogenetic reconstructions suggested that EFN-bearing species are monophyletic, whereas the evolution of domatia correlated with leaf production strategy (deciduous or evergreen) and climate. Domatia were also more common in the EFN clade, suggesting that the two traits may jointly mediate ecological interactions. This result was further investigated in a common-garden survey, where plants with domatia and EFNs on the leaf blade had more mutualistic mites than plants with other trait combinations, and in manipulative field experiments, where the traits additively increased mutualist abundance. Taken together, our results suggest that mutualistic traits in Viburnum are not ecologically independent, as they work in concert to attract and retain mutualists, and their long-term evolution may be influenced by complex interactions among multiple traits, mutualists, and geography.     

毛茛科铁线莲属的叶表皮特征及其系统学意义(英文)

用光学显微镜和扫描电镜 ,较全面地观察了铁线莲属 (Clematis)植物和邻近类群的叶表皮特征 ,并对其系统学意义进行了讨论。在光学显微镜下 ,观察了铁线莲属 12组 6 3种 5变种及其同族邻近类群 3属 9种 ;在扫描电镜下 ,观察了铁线莲属 7组 2 7种 2变种及其邻近类群 2属 6种植物的成熟叶表皮细胞及气孔器的特征。在光学显微镜下可见 :叶表皮细胞形状 (表面观 )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 ,少数种的上表皮也有分布 ,其类型均为无规则型。在扫描电镜下可见 :叶片通常具有条纹或有颗粒状或鳞片状附属物 ,气孔器外拱盖内缘通常为浅波状或近平滑。在黄花铁线莲组 (sect.Meclatis (Spach)Tamura)和灌木铁线莲组 (sect.FruticellaTamura)中 ,叶表皮具有双面气孔器的特征对于组下亚组的划分有一定意义。并且叶表皮特征对于一些种的分类学处理也有一定的价值 ,同时也为属下组间的系统进化关系提供了一些线索     

Leaf domatia reduce intraguild predation among predatory mites

1. Although theory suggests that intraguild predation destabilises food webs and may result in exclusion of species, empirical observations of food webs reveal that it is a common interaction. It has been proposed that habitat structure reduces the interaction strength of intraguild predation, thus facilitating the coexistence of species. 2. This was tested using acarodomatia, tiny structures on plant leaves, and predatory mites, which usually reside in these domatia. Sweet pepper plants (Capsicum annuum L.) were used, which possess domatia consisting of tufts of hair, and coffee plants (Coffea arabica L.) with pit‐shaped domatia. 3. On sweet pepper, the predatory mites Neoseiulus cucumeris Oudemans and Iphiseius degenerans Berl. feed on each other's juveniles. Larvae of each of the species were therefore used as intraguild prey with adult females of the other species as intraguild predators. On coffee, a similar set‐up was used, with larvae and adult females of Amblyseius herbicolus Chant and Iphiseiodes zuluagai Denmark & Muma as intraguild prey and intraguild predators, respectively. 4. Domatia on detached, isolated sweet pepper and coffee leaves were either closed with glue or left open, after which larvae and adult predators were released. As a control, larvae were released on leaves with open or closed domatia without an adult predator. 5. Survival of larvae was high in the absence of the adult (intraguild) predator. In the presence of the intraguild predator, survival was significantly higher on leaves with open domatia than on leaves with closed domatia. 6. This shows that even such tiny structures as plant domatia may significantly affect the interaction strength of intraguild predation.     

鄂尔多斯高原锦鸡儿属植物叶表皮特征及生态适应性

在光学显微镜和扫描电镜下,对鄂尔多斯高原上从东到西分布的8种锦鸡儿属(Caragana)植物的叶表皮特征进行了观察,发现该属植物的叶表皮细胞形状有多边形和不规则形两种类型;气孔轴式类型主要是不定式。旱中生的秦晋锦鸡儿(C. purdomii)的气孔密度及气孔指数最高,其它7种旱生种类(甘蒙锦鸡儿(C. opulens)、中间锦鸡儿(C. intermedia)、柠条锦鸡儿(C. korshinskii)、荒漠锦鸡儿(C. roborovskyi)、狭叶锦鸡儿(C. stenophylla)、藏锦鸡儿(C. tibetica)和短脚锦鸡儿(C. brachypoda))的偏低。在同种锦鸡儿中,中间锦鸡儿、狭叶锦鸡儿、藏锦鸡儿的气孔密度随环境干旱程度的增加而增加,柠条锦鸡儿的气孔密度则减少;狭叶锦鸡儿、藏锦鸡儿的气孔指数随环境干旱程度的增加表现为上升,中间锦鸡儿、柠条锦鸡儿的气孔指数表现为下降。中间锦鸡儿、柠条锦鸡儿、藏锦鸡儿和荒漠锦鸡儿的表皮均密被表皮毛,是对干旱环境的适应特征。旱生种类比中生种类的表皮特性具有更高的可塑性,在半荒漠地带能够成为建群种的狭叶锦鸡儿和藏锦鸡儿的可塑性指数最高。在扫描电镜下,气孔位置有两类:气孔与表皮细胞齐平、气孔陷于表皮细胞之下。表皮上有角质层和蜡质;表皮毛为单细胞非腺毛。     

Collagen deposition on a preformed grid

Appearance of collagen fibrils in the cuticle was seen by electron microscopy to be preceded by fonnation of a finely filamentous matrix material. At first, the fine filaments of the matrix are unorganized. However, signs of orthogonal ordering soon appear in the most superficial portion of the cuticle, and subsequently appear more basally and closer to the underlying epidermis. Meanwhile, fibrils of different staining properties and identifiable as collagen begin to be deposited in the superficial portion of the cuticle, the same region which first showed organized fine filaments. Then, like the fine filaments before them, the collagen fibrils polymerize more basally. Collagen appears to polymerize on the preformed skeleton of fine filaments as though the fine filaments caused the collagen to assemble. Neither the polymerization nor ordering of collagen fibrils seems to require direct cellular intervention but occur first in that portion of the cuticle which is furthest away from the underlying epidermis. The fine filaments may be self ordering, extracellular macromolecules which in turn determine the polymerization of collagen fibrils.     

Behavioral and population consequences of acarodomatia in grapes on phytoseiid mites (Mesostigmata) and implications for plant breeding

We examined the influence of acarodomatia in the riverbank grape Vitis riparia Michaux (Vitaceae) on the distribution and abundance of predatory mites (Phytoseiidae) and their interactions with herbivorous mites. Acarodomatia are tufts of nonglandular trichomes or pits located in major leaf vein axes of many species of woody perennial plants and are often occupied by predatory and mycophagous mites. In common garden plantings of different accessions of V. riparia we found a significant positive relationship between size of domatia and the abundance of naturally occurring predatory mites. Behavior of adult predatory mites may explain this positive association, in part. In separate laboratory experiments, gravid females of Typhlodromus pyri Scheuten and Amblyseius andersoni Chant spent more time and deposited more eggs on half of a V. riparia leaf with accessible domatia versus the other half in which access to domatia was blocked with pruning tar. Domatia also had population consequences. In an outdoor experiment using potted grapevines, population size of T. pyri and A. andersoni mites was greater on V. riparia with open domatia compared to V. riparia in which domatia were blocked with pruning tar. Population size of predatory mites was also greater on V. riparia with domatia than on Vitis vinifera L., whether their axils were blocked or not. Since V. vinifera have very small domatia, these results indicate that the presence of domatia is important, not just access to vein axils. Elevated predatory mite populations in response to domatia, however, did not translate into differences in the abundance of European red mite Panonychus ulmi (Koch), an important pest of grapes.Overall, these results indicate that domatia in uncultivated V. riparia promote higher densities of some species of generalist phytoseiid mites. However, domatia are small in most cultivated grapes. We crossed females and males of V. riparia that varied in domatia size and reared their offspring and found that average domatia size in the parents was highly correlated (r² = 0.77, slope = 0.55) with average domatia size in offspring (high narrow-sense heritability). Given that V. riparia possesses many other desirable agronomic traits, this result suggests it should be practical to breed for well-developed domatia in cultivated accessions.     

Changes of Leaf Morphological, Anatomical Structure and Carbon Isotope Ratio with the Height of the Wangtian Tree （Parashorea chinensis） in Xishuangbanna, China

Leaf morphological and anatomical structure and carbon isotope ratio （δ^13C） change with increasing tree height. To determine how tree height affects leaf characteristics, we measured the leaf area, specific leaf mass （ratio of leaf mass to leaf area [LMA]）, thickness of the total leaf, cuticle, epidermis, palisade and sponge mesophyll, stomata traits and δ^13C at different heights of Parashorea chinensis with methods of light and scanning electron microscopy （SEM） and isotope-ratio mass spectrometry. The correlation and stepwise regression between tree height and leaf structure traits were carried out with SPSS software. The results showed that leaf structures and δ^13C differed significantly along the tree height gradient. The leaf area, thickness of sponge mesophyll and size of stomata decreased with increasing height, whereas the thickness of lamina, palisade mesophyll, epidermis, and cuticle, ratios of palisade to spongy thickness, density of stomata and vascular bundles, LMA and δ^13C increased with tree height. Tree height showed a significant relationship with all leaf indices and the most significant relationship was with epidermis thickness, leaf area, cuticle thickness, δ^13C. The δ^13C value showed a significantly positive relationship with LMA （R = 0.934）. Our results supported the hypothesis that the leaf structures exhibited more xeromorphic characteristics with the increasing gradient of tree height.     

中国棱子芹属植物叶表皮微形态特征及其系统学研究

采用光学显微镜和扫描电镜对棱子芹属(Pleurospermum)11种植物的叶表皮微形态特征进行了观察研究.结果表明:(1)显微观察显示,气孔器普遍存在于叶的下表皮.少数种的上表皮也有分布,多为无规则型;叶表皮细胞形状为多边型或不规则形.垂周壁式样可区分为近平直、平直-弓形、浅波状和波状.(2)扫描电镜观察显示,叶表皮气孔器外拱盖内缘为近平滑、浅波状或波状;角质膜条纹状,有的条纹隆起,有的条纹上附有颗粒和晶簇.(3)气孔器的分布、气孔器外拱盖内缘形态以及角质膜等特征对该属部分种的区分有一定的参考价值.(4)结合外部形态特征,推测康定棱子芹应作为种一级分类群.松潘棱子芹与宝兴棱子芹的亲缘关系比较近.     

鹿药属植物叶表皮特征及其系统学意义

采用光学显微镜和扫描电镜对鹿药属12种植物的叶表皮进行了观察,首次报道了12种鹿药属(Sm ilaci-na)植物叶表皮的微形态特征。结果表明:气孔器普遍存在于叶的下表皮,少数种的上表皮也有分布,均为不规则形。叶表皮细胞形状为多边形或不规则形,垂周壁式样可区分为近平直、浅波状和波状。在扫描电镜下,叶表皮气孔器外拱盖内缘为近平滑、浅波状或波状;角质膜条纹状,有的条纹隆起,有的条纹上附有颗粒和晶簇。气孔器的分布、气孔器外拱盖内缘形态以及角质膜等特征对该属部分种的区分具有一定的参考价值。     

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.