Optical and anatomical characteristics of bracts from the Chinese "glasshouse" plant, Rheum alexandrae Batalin (Polygonaceae), in Yunnan, China

Bracts that lacked chlorophyll were compared with rosette leaves on the Chinese glasshouse plant Rheum alexandrae Batalin. The structures were analyzed anatomically and with photospectrometry. Histological features were significantly different between the bracts and the rosette leaves. Epidermal pavement cells and palisade cells were larger in the bracts than in the rosette leaves, but the ratio of the intercellular spaces to the cells in the palisade layer was the same in the two structures. Absorption spectrum analyses of the half-translucent bracts showed that, despite their thinner structure, they absorbed ultraviolet (UV) radiation more efficiently than did the green rosette leaves. The characteristics of the R. alexandrae bracts were different from those of the Himalayan glasshouse plant Rheum nobile. These results suggest that R. alexandrae evolved bracts as an adaptive strategy to protect the inflorescence from UV radiation. Received: August 22, 2001 / Accepted: October 15, 2001     

Selective light transmittance of translucent bracts in the Himalayan giant glasshouse plant Rheum nobile Hook. f. & Thomson (Polygonaceae)

Translucent bract transmittance of ultraviolet (UV) to infrared (IR) radiation (between 320 and 800 nm) and leaf anatomy were examined in a glasshouse plant, Rheum nobile Hook. f. & Thomson (Polygonaceae) to assess the function of avoiding injury by UV radiation while keeping the inflorescence warm by photosynthetically active (PA) and IR radiation. Although the translucent bracts and rosulate leaves transmitted little UV radiation, the former always transmit more PA and IR radiation. Additionally, the bracts transmit much more scattered solar radiation than direct radiation. The bracts are also anatomically different from the rosulate leaves. They have two or three layers of mesophyll cells with neither palisade nor spongy parenchymatous cells; in addition, the uppermost layer of mesophyll and the epidermis stain easily, and both are thought to play a role in attenuating UV radiation. The leaf epidermis of many land plants has UV absorbing pigments such as flavonoids, which absorb almost all UV radiation. Thus the role of the bracts of R. nobile is to protect the reproductive organs by absorbing UV radiation and to keep them warm by transmitting PA and IR radiation. The bracts are believed to have adapted function and form to the environment, in particular, to the weather conditions of the eastern Himalaya.     

不同色彩珙桐叶片和苞片解剖结构及色素含量比较研究

以生长于同一生境下的粉红珙桐(粉红色叶片、苞片)与普通珙桐(绿色叶片、白色苞片)为试材,对比两种色彩珙桐叶片/苞片解剖结构和色素含量的差异,以揭示珙桐色彩转变的规律。结果显示：(1)两种珙桐叶片均属于异面叶类型,栅栏组织由一层长柱形细胞整齐排列而成,海绵组织排列疏松,部分粉红叶片的上表皮细胞向外凸起,绿叶无此现象;粉红叶片的总厚度及其表皮角质层、栅栏组织和海绵组织厚度都高于绿叶,而表皮较薄。(2)两种珙桐苞片均无栅栏组织和海绵组织的分化,粉红苞片上表皮细胞明显隆起,上表皮角质层增厚,而下表皮变薄。(3)粉红叶片的类黄酮、花色苷含量分别是绿色叶片的1.52倍、3.67倍,两者的光合色素含量无显著差异,但粉红叶片的叶绿素a/b值比绿色叶低很多;粉红苞片花色苷含量显著高于白色苞片,而两者类黄酮含量差异不大。研究表明,花色苷是珙桐叶片和苞片色彩转红的直接因素,类黄酮有助于叶片呈红色;粉红珙桐叶片/苞片的解剖结构发生了一定变化,对光能的利用效率更高,对阴湿环境的适应性增强。     

Physiological characterization of photosynthesis,chloroplast ultrastructure,and nutrient content in bracts and rosette leaves from <Emphasis Type="Italic">Glaucium flavum</Emphasis>

Glaucium flavum is a biennial plant that bears a rosette of leaves, producing a flower stalk, bracteate monochasium, in its second year. The aims of this work were both to investigate the contribution of bracts to gas-exchange activities in this species and to compare this contribution to that of rosette leaves. In addition, we investigated the extent to which its responses can be explained by chloroplast ultrastructure, as well as the possible role of nutrient concentrations in the physiological responses of both leaf types. Gas exchange and plant characteristics regarding chlorophyll fluorescence were examined in a field experiment; we also determined leaf relative water content, tissue concentrations of photosynthetic pigments, chloroplast ultrastructure and nutrient contents. Although bracts indeed contributed to gas-exchange activities of G. flavum, rosette leaves showed higher values of net photosynthetic rate and stomatal conductance to CO₂ for photosynthetic photon flux density above 200 μmol m⁻² s⁻¹. The incongruities in photosynthetic rates between bracts and leaves may be explained by the bigger chloroplasts of rosette leaves, which results in a larger membrane surface area. This agrees with the higher pigment concentrations and quantum efficiency of photosystem II values recorded as well for rosette leaves. On the other hand, bracts showed higher sodium concentrations, which could be a mechanism for salt tolerance of G. flavum.     

Molecular phylogeny, recent radiation and evolution of gross morphology of the rhubarb genus Rheum (Polygonaceae) inferred from chloroplast DNA trnL-F sequences

BACKGROUND AND AIMS: Rheum, a highly diversified genus with about 60 species, is mainly confined to the mountainous and desert regions of the Qinghai-Tibetan plateau and adjacent areas. This genus represents a good example of the extensive diversification of the temperate genera in the Qinghai-Tibetan plateau, in which the forces to drive diversification remain unknown. To date, the infrageneric classification of Rheum has been mainly based on morphological characters. However, it may have been subject to convergent evolution under habitat pressure, and the systematic position of some sections are unclear, especially Sect. Globulosa, which has globular inflorescences, and Sect. Nobilia, which has semi-translucent bracts. Recent palynological research has found substantial contradictions between exine patterns and the current classification of Rheum. Two specific objectives of this research were (1) to evaluate possible relationships of some ambiguous sections with a unique morphology, and (2) to examine possible occurrence of the radiative speciation with low genetic divergence across the total genus and the correlation between the extensive diversification time of Rheum and past geographical events, especially the recent large-scale uplifts of the Qinghai-Tibetan Plateau. METHODS: The chloroplast DNA trnL-F region of 29 individuals representing 26 species of Rheum, belonging to seven out of eight sections, was sequenced and compared. The phylogenetic relationships were further constructed based on the sequences obtained. KEY RESULTS: Despite the highly diversified morphology, the genetic variation in this DNA fragment is relatively low. The molecular phylogeny is highly inconsistent with gross morphology, pollen exine patterns and traditional classifications, except for identifying all samples of Sect. Palmata, three species of Sect. Spiciformia and a few species of Sect. Rheum as corresponding monophyletic groups. The monotypic Sect. Globulosa showed a tentative position within the clade comprising five species of Sect. Rheum. All of the analyses revealed the paraphyly of R. nobile and R. alexandrae, the only two species of Sect. Nobilia circumscribed by the possession of large bracts. The crude calibration of lineages based on trnL-F sequence differentiation implied an extensive diversification of Rheum within approx. 7 million years. CONCLUSIONS: Based on these results, it is suggested that the rich geological and ecological diversity caused by the recent large-scale uplifts of the Qinghai-Tibetan Plateau since the late Tertiary, coupled with the oscillating climate of the Quaternary stage, might have promoted rapid speciation in small and isolated populations, as well as allowing the fixation of unique or rare morphological characters in Rheum. Such a rapid radiation, combined with introgressive hybridization and reticulate evolution, may have caused the transfer of cpDNA haplotypes between morphologically dissimilar species, and might account for the inconsistency between morphological classification and molecular phylogeny reported here.     

华东椴花序苞片和叶片发育及结构特征比较研究

花序苞片是椴树属最典型的特征,但它与其他器官的演化关系尚不明确。本文对华东椴（Tilia japonica Simonk.）花序苞片和叶片在生长发育上的关联性进行分析,并对其表面微形态特征和内部解剖结构进行详细比较。结果显示,华东椴苞片和叶片在生长发育过程中表现出非常相似的规律性,二者在苞片幼期阶段均生长迅速、中期阶段生长变缓、成熟阶段再有小幅生长。对苞片与叶片形态性状的相关性分析显示,苞片长、苞片宽、叶长、叶宽的相关性均极显著。在表面微形态特征上,苞片和叶片总体上非常相似;在解剖特征上,苞片没有栅栏组织和海绵组织的分化,叶片有明显的分化。结合椴树属苞片的化石记录,我们认为椴树属苞片最早由叶片特化而来,是一种叶状结构,二者在表皮微形态特征及解剖结构上的差异,主要是在演化过程中为了适应各自不同功能而逐渐产生的。     

Anatomical considerations related to photosynthesis in cotton (Gossypium hirsutum L.) leaves, bracts, and the capsule wall

Light and electron microscopy was used to relate histologicaland ultrastructural differences of cotton (Gossypium hirsutumL.) leaves, bracts, and capsule walls to their different photosyntheticactivities. Light microscopy revealed that the leaf thicknesswas approximately 152µm, had a well-defined internal organizationwith elongated palisade mesophyll cells and loosely packed spongymesophyll cells. In contrast, the bract was thinner (111 µm),lacked a defined palisade layer, and was largely composed ofinternal air spaces. The capsule wall was very thick (1013µm)and composed of numerous tightly packed, paren-chymatous corticalcells with little or no intercellular air space. Chloroplastswith well-defined granal stacks and extensive stroma lamellaewere observed in each of these three tissues, however, theirdensity was always greater in the palisade cells of the leafcompared to spongy mesophyll cells of the bract and the parenchymatouscells of the capsule wall. The low rates of photosynthesis inthe bracts and the capsule wall were associated with the internalorganization of these tissues. Key words: Cotton, photosynthesis, anatomy, cuticle, tissues     

蒙古莸叶片解剖结构的地理种源变异及其对环境变化响应的意义

长期受到生长环境影响而形成的遗传变异对植物生长发育有着显著的影响。叶片是植物对环境变化最敏感的器官, 了解叶片解剖结构在不同环境中产生的适应性变异是探索植物对环境适应的基础。同质园试验是研究遗传与环境因素对植物生长代谢等影响的一种有效方法, 该研究利用同质园试验排除了环境梯度的影响, 通过常规石蜡切片、多重比较、相关性分析、一般线性模型分析等方法, 对7个不同种源地的蒙古莸(Caryopteris mongholica)叶片解剖结构及其影响因素进行了定量比较。结果表明, 7个种源地的蒙古莸叶片均为等面叶, 无海绵组织分化, 其上表皮细胞较下表皮细胞厚, 上栅栏组织较下栅栏组织厚; 叶片各解剖结构参数间存在显著的自相关性, 不同种源叶片解剖结构存在显著差异: 随种源地年平均气温升高, 叶厚度、栅栏组织厚度呈增大趋势, 其中, 最西南部的阿左旗种源蒙古莸叶片的上下栅栏组织、叶厚度及叶片结构紧密度值均最大, 表现出明显的抗旱特征。种源地经纬度、气温、降水等对解剖结构指标有显著的影响, 其解释程度为34.09%-81.43%。同质园试验说明, 种源地气候差异驱动的遗传变异是引起不同种源叶片解剖结构差异的重要因素。     

Multifunctional bracts enhance plant fitness during flowering and seed development in Rheum nobile (Polygonaceae), a giant herb endemic to the high Himalayas

Specialized bracts are thought to be important for the successful reproduction of some plants and are regarded as adaptations to diverse driving forces. However, few empirical studies have quantified the adaptive significance of bracts within a cost–benefit framework. We explored the adaptive significance of large and showy bracts for reproduction in Rheum nobile, a giant herb endemic to the high Himalayas. We examined whether the bracts enhance reproductive success during flowering and seed development. Bracts increased flower and fruit temperature on sunny days, greatly decreased the intensity of ultraviolet-B (UV-B) radiation reaching flowers and fruits, and prevented pollen grains being washed away by rain. Experiments indicated that high temperature could promote pollen germination, while pollen grains exposed to rain and UV-B radiation at ambient levels were seriously damaged. Furthermore, bract removal decreased the number of pollinators visiting flowers. When bracts were removed before or after flowering, fecundity and progeny quality were adversely affected, but seed predation by larvae of pollinators decreased. A cost–benefit analysis demonstrated that the cost of bracts, i.e., increased seed predation, is modest. Our results suggest that the bracts of R. nobile promote pollen germination, protect pollen grains from rain and intense UV-B radiation, enhance pollinator visitation during flowering, and facilitate the development of fertilized ovules during seed development. We conclude that multifunctional bracts of R. nobile are an effective adaptive strategy in alpine environments and might have been selected for because of abiotic environmental conditions as well as for enhancing pollination success.     

Flavonoids in translucent bracts of the Himalayan<Emphasis Type="Italic"> Rheum nobile</Emphasis> (Polygonaceae) as ultraviolet shields

UV-absorbing substances were isolated from the translucent bracts of Rheum nobile, which grows in the alpine zone of the eastern Himalayas. Nine kinds of the UV-absorbing substances were found by high performance liquid chromatography (HPLC) and paper chromatography (PC) surveys. All of the five major compounds are flavonoids, and were identified as quercetin 3-O-glucoside, quercetin 3-O-galactoside, quercetin 3-O-rutinoside, quercetin 3-O-arabinoside and quercetin 3-O-[6-(3-hydroxy-3-methylglutaroyl)-glucoside] by UV, ¹H and ¹³C NMR, mass spectra, and acid hydrolysis of the original glycosides, and direct PC and HPLC comparisons with authentic specimens. The four minor compounds were characterised as quercetin itself, quercetin 7-O-glycoside, kaempferol glycoside and feruloyl ester. Of those compounds, quercetin 3-O-[6-(3-hydroxy-3-methylglutaroyl)-glucoside] was found in nature for the first time. The translucent bracts of R. nobile accumulate a substantial quantity of flavonoids (3.3–5 mg per g dry material for the major compounds). Moreover, it was clarified by quantitative HPLC survey that much more of the UV-absorbing substances is present in the bracts than in rosulate leaves. Although the flavonoid compounds have been presumed to be the important UV shields in higher plants, there has been little characterisation of these compounds. In this paper, the UV-absorbing substances of the Himalayan R. nobile were characterised as flavonol glycosides based on quercetin.     

Red : far-red light ratio and UV-B radiation: their effects on leaf phenolics and growth of silver birch seedlings

The natural variation in quantity and quality of light modifies plant morphology, growth rate and concentration of biochemicals. The aim of two growth‐room experiments was to study the combined effects of red (R) and far‐red (FR) light and ultraviolet‐B (UV‐B) radiation on the concentrations of leaf phenolics and growth and morphology of silver birch (Betula pendula Roth) seedlings. Analysis by high‐performance liquid chromatography showed that the leaves exposed to supplemental FR relative to R contained higher concentrations of total chlorogenic acids and a cinnamic acid derivative than the leaves treated with supplemental R relative to FR. In contrast, concentration of a flavonoid, quercetin 3‐galactoside, was higher in the R + UV‐B leaves than in the FR + UV‐B leaves. The UV‐B induced production of kaempferols, chlorogenic acids and most quercetins were not modified by the R : FR ratio. Growth measurements showed that the leaf petioles and stems of FR seedlings were clearly longer than those of R seedlings, but leaf area was reduced by UV‐B radiation. Results of these experiments show that exposure of silver birch seedlings to supplemental FR compared to R leads to fast elongation growth and accumulation of phenolic acids in the leaves.     

云南秋海棠属植物叶片横切面比较解剖研究

报道30种主产于云南的秋海棠属植物叶片的横切面解剖构造特征。采用常规石蜡切片法切片观察,结果表明:云南秋海棠属植物叶片薄、横切面均为异面叶、呈典型的阴叶结构,叶肉组织虽有栅栏组织和海绵组织的分化,但栅栏组织不发达,占叶肉组织的比例较小。表皮多为单表皮,极稀复表皮,表皮毛均由多细胞组成。气孔集中于下表皮,孔下室或下陷气孔特大、通气组织极发达;角质层形状多样,呈均匀增厚、瘤状和片状突起;叶绿体椭球形、数多、个体大,主要分布于叶肉组织,集中于栅栏组织。解剖构造特征在各分类组内呈现不完全一致性,而在相同茎的形态类型中有些较一致的特征,在不同种间解剖特征各有差别;根状茎和直立茎类型种类的横切面组织结构表现为表皮细胞壁外的角质层薄、栅栏组织与叶肉组织厚度比例较小等弱光照、湿生等适应性较弱的特征。球茎类型的种类表现为角质层较厚、栅栏叶肉组织厚度比例较大等适应略为干燥和较强光照的特征。     

藏东南色季拉山薄毛海绵杜鹃叶解剖结构特征与环境适应性

以藏东南色季拉山9个不同海拔梯度的薄毛海绵杜鹃(Rhododendron aganniphum var.schizopeplum)叶片为试验材料,采用石蜡切片技术,测定10项叶片解剖结构指标,应用叶片解剖结构指标的可塑性指数和相关性分析方法探索薄毛海绵杜鹃对藏东南色季拉山强紫外辐射和高寒环境的适应性。结果表明:(1)薄毛海绵杜鹃叶片为异面叶,上表皮有明显的角质层,下表皮有表皮毛,栅栏组织细胞2～3层。(2)随着海拔的升高,叶片角质层厚度、上下表皮厚度、栅栏组织厚度、海绵组织厚度、叶片厚度呈现明显增大趋势,而组织结构紧密度和疏松度变化不显著,主脉突起度呈现下降趋势。(3)各项叶片解剖结构指标的可塑性指数显示,薄毛海绵杜鹃在解剖结构上表现出较小的可塑性,对外界环境的适应能力较弱。(4)依据薄毛海绵杜鹃各项叶片解剖结构指标的相关性分析结果,除海绵组织厚度与栅栏组织厚度、下表皮厚度与上角质层厚度之间相关性不显著外,其余各指标之间均呈显著相关关系,且叶片的解剖结构指标方面也存在明显的协同进化现象。研究发现,藏东南色季拉山薄毛海绵杜鹃通过增加叶片角质层厚度、表皮厚度和叶肉厚度等解剖结构指标的方式增强对外界极端环境的适应能力,从而有利于其在恶劣的高山生境下生存繁衍,使该物种成为生态位理论中的广幅种。     

补增UV-B辐射对香蕉叶片光合作用和叶氮在光合碳循环组分中分配的影响

补增UV－B辐射的香蕉叶片光下呼吸速率（Rd））和不包括光下呼吸的CO2补偿点（г*）,分别为0.33μmol·m-2·s-1和46.5μl·L-1,较对照植株分别高5.6％和10.0％。在较高CO2浓度（>340μl·L-1）条件下的An/θp关系最初直线部分斜率,即表观量子产率(α-A)为0.023±0.007,而补增UV-B辐射处理的植株则降低13.0％,光能转换效率（δ）亦降低28.6％,表明UV-B辐射明显降低αA和δ。在高θp（1100μmol·m-2·s-1）和Ci<200μl·L-1条件下,对照植株的An/Ci关系为An=0.028Ci+1.44,补增UV-B辐射处理的植株则为An=0.021Ci+1.01,UV-B辐射降低羧化限制速率。最大羧化速率（Vcmax）和电子传导速率的光饱和值（Jmax）亦较低,补增UV-B辐射的叶片,叶氮在Rubisco的分配系数（PR）和叶氮在生物力能学组分的分配系数（PB）分别较对照低8.1％和3.0％,叶氮分配到类囊体膜捕光色素蛋白组分的则略见增高,UV-B辐射降低叶氮在光合循环组分的分配。     

Effect of UV-B Radiation on Leaf Optical Properties Measured with Fibre Optics

Changes in the internal light microenvironment in leaves ofplants of Brassica campestris L. cv. Emma, B. carinata L., andMedicago saliva L. cv. Armour in response to exposure to UV-B(UV-B, 280–320 nm) radiation were measured using a fibreopticmicroprobe. Plants were exposed for 2 weeks either to high visiblelight or to supplemental ultraviolet-B radiation. The spectral regime (400–700 nm; PAR) was measured eithermidway through the leaf palisade or the spongy mesophyll. Afterexposure to UV-B radiation leaves of Brassica campesiris attenuatedtransmitted light more than the controls. At the same time bothforward and back scattered light increased in the palisade andspongy mesophylls. In contrast, UV-treatment of Medicago salivaleaves increased light transmission into the palisade, whilethe back scattered component showed little change. Leaves ofcariiwla showed little change in response to UV. Other responsesto UV-B radiation included increases in leaf thickness, decreasedtotal chlorophyll content, and changes in UV-B screening pigmentsand chlorophyll fluorescence induction kinetics. Brassica campestriswas most sensitive to exposure to enhanced levels of UV-B radiation,whereas leaves of B. carinata were the least sensitive. Ourdata indicate that exposure to UV-B radiation altered the lightmicroenvironment within leaves of the species different ways.These changes appeared to be caused by alterations in pigmentcontent and leaf anatomy. In turn, the altered distributionof PAR within the leaf could influence photosynthesis. Key words: Brassica campestris, Brassica carinata, fibre optics, light scattering, Medicago saliva, optical properties, ozone depletion, photosynthesis, ultraviolet radiation     

海桑和无瓣海桑叶片结构的比较研究

为了探讨外来植物无瓣海桑的潜在危害,采用石蜡切片法对海桑(Sonneratia caseolaris Engl.)、无瓣海桑(S.apetala B.Ham)叶片进行了解剖学研究。实验结果显示,两种植物的叶片均为等面叶;中脉维管束为周韧维管束;具4级侧脉,第1级侧脉为半周韧维管束;成熟叶片叶肉组织具发达的贮水薄壁细胞,具含单宁成分的薄壁细胞,具晶体细胞和石细胞;盐腺由表皮细胞发育而成,可分为3个发育阶段。作为外来植物的无瓣海桑,其中脉维管束具微弱形成层,叶脉维管组织比海桑更发达;贮藏组织中含单宁细胞、晶体细胞较多;栅栏组织含叶绿体多于海桑等特点,使其比海桑对环境具有更大的适应性。因此,无瓣海桑有可能成为入侵植物。     

Euphrasia brevilabris Y. F. Wang,Y. S. Lian & G. Z. Du,a new species of the Scrophulariaceae from the Qinghai-Tibetan Plateau,China

Euphrasia brevilabris Y. F. Wang, Y. S. Lian & G. Z. Du, a new species of the Scrophulariaceae from Gannan grassland of Gansu belonging to the east edge of the Qinghai-Tibetan Plateau of China, is described and illustrated. It is related to E. regelii Wettst. by having dense tentacle on the leaves, bracts and calyces, but differs from plant shorter, 3–7 cm high, without ramification, leaves smaller, 3–5 mm long, corolla yellow, the back 3–5 mm long, and labium shorter than muffle. It is also similar to E. pectinata Ten. by bracts obviously bigger than vegetative leaves, but differs from plant shorter, leaves smaller, having dense tentacle on the leaves, bracts and calyces, corolla yellow, labium shorter than muffle.