Association between photosynthesis and contrasting features of minor veins in leaves of summer annuals loading phloem via symplastic versus apoplastic routes

Foliar vascular anatomy and photosynthesis were evaluated for a number of summer annual species that either load sugars into the phloem via a symplastic route (Cucumis sativus L. cv. Straight Eight; Cucurbita pepo L. cv. Italian Zucchini Romanesco; Citrullus lanatus L. cv. Faerie Hybrid; Cucurbita pepo L. cv. Autumn Gold) or an apoplastic route (Nicotiana tabacum L.; Solanum lycopersicum L. cv. Brandywine; Gossypium hirsutum L.; Helianthus annuus L. cv. Soraya), as well as winter annual apoplastic loaders (Spinacia oleracea L. cv. Giant Nobel; Arabidopsis thaliana (L.) Heynhold Col‐0, Swedish and Italian ecotypes). For all summer annuals, minor vein cross‐sectional xylem area and tracheid number as well as the ratio of phloem loading cells to phloem sieve elements, each when normalized for foliar vein density (VD), was correlated with photosynthesis. These links presumably reflect (1) the xylem's role in providing water to meet foliar transpirational demand supporting photosynthesis and (2) the importance of the driving force of phloem loading as well as the cross‐sectional area for phloem sap flux to match foliar photosynthate production. While photosynthesis correlated with the product of VD and cross‐sectional phloem cell area among symplastic loaders, photosynthesis correlated with the product of VD and phloem cell number per vein among summer annual apoplastic loaders. Phloem cell size has thus apparently been a target of selection among symplastic loaders (where loading depends on enzyme concentration within loading cells) versus phloem cell number among apoplastic loaders (where loading depends on membrane transporter numbers).     

Leaf architectural,vascular and photosynthetic acclimation to temperature in two biennials

Acclimation of leaf features to growth temperature was investigated in two biennials (whose life cycle spans summer and winter seasons) using different mechanisms of sugar loading into exporting conduits, Verbascum phoeniceum (employs sugar‐synthesizing enzymes driving symplastic loading through plasmodesmatal wall pores of phloem cells) and Malva neglecta (likely apoplastic loader transporting sugar via membrane transport proteins of phloem cells). In both species, acclimation to lower temperature involved greater maximal photosynthesis rates and vein density per leaf area in close correlation with modification of minor vein cellular features. While the symplastically loading biennial exhibited adjustments in the size of minor leaf vein cells (consistent with adjustment of the level of sugar‐synthesizing enzymes), the putative apoplastic biennial exhibited adjustments in the number of cells (consistent with adjustment of cell membrane area for transporter placement). This upregulation of morphological and anatomical features at lower growth temperature likely contributes to the success of both the species during the winter. Furthermore, while acclimation to low temperature involved greater leaf mass per area in both species, this resulted from greater leaf thickness in V. phoeniceum vs a greater number of mesophyll cells per leaf area in M. neglecta. Both types of adjustments presumably accommodate more chloroplasts per leaf area contributing to photosynthesis. Both biennials exhibited high foliar vein densities (particularly the solar‐tracking M. neglecta), which should aid both sugar export from and delivery of water to the leaves.     

Relationship between photosynthetic rate,water use and leaf structure in desert annual and perennial forbs differing in their growth

Specific leaf area (SLA) is a key trait to screen plants for ecological performance and productivity; however, the relationship between SLA and photosynthesis is not always up-scalable to growth when comparing multiple species with different life cycles. We explored leaf anatomy in annual and perennial species of Physaria, and related it to photosynthesis and water loss. The annual Physaria gracilis had higher SLA, thinner leaves, and lower investment in protective tissues, than perennial P. mendocina. Physaria angustifolia (annual), and P. pinetorum (perennial) showed intermediate values. Both perennials had a thicker palisade and high photosynthesis, relative to annuals. The larger leaf veins of perennials should allow high water availability to the mesophyll. The thicker palisade should determine high resistance to water flow and help explain their high water-use efficiency. These leaf functions reflect the construction of long-lived leaves that efficiently use resources under environmental limitations of arid environments.

     

沙棘雌雄株叶片解剖结构比较研究

为了研究沙棘雌、雄株叶片的第二性征,本文采用石蜡切片法观察了沙棘雌、雄株叶片结构的差异。结果表明：（1）沙棘雌、雄株叶片均由表皮、叶肉和叶脉3部分组成,表皮均由1层细胞构成,表皮毛发达,上表皮有拟泡状细胞;叶肉栅栏组织与海绵组织分化明显。（2）雌株上表皮具更多的拟泡状细胞,其主脉韧皮部薄壁细胞及其下方的一些薄壁细胞含较多的后含物,下表皮的表皮毛更浓密;而雄株的叶片厚度、叶片上表皮厚度、栅栏组织厚度、栅栏组织厚度/海绵组织厚度均显著大于雌株,且其主脉维管束更发达。结果表明,沙棘雌雄株叶片解剖结构存在明显差异,这些差异是第二性征的表现,也是沙棘长期进化中形成的稳健的适应策略,可能有利于该物种的繁衍。     

Sun-shade adaptability of the red mangrove,Rhizophora mangle (Rhizophoraceae): Changes through ontogeny at several levels of biological organization

Rhizophora mangle L., the predominant neotropical mangrove species, occupies a gradient from low intertidal swamp margins with high insolation, to shaded sites at highest high water. Across a light gradient, R. mangle shows properties of both “light-demanding” and “shade-tolerant” species, and defies designation according to existing successional paradigms for rain forest trees. The mode and magnitude of its adaptability to light also change through ontogeny as it grows into the canopy. We characterized and compared phenotypic flexibility of R. mangle seedlings, saplings, and tree modules across changing light environments, from the level of leaf anatomy and photosynthesis, through stem and whole-plant architecture. We also examined growth and mortality differences among sun and shade populations of seedlings over 3 yr. Sun and shade seedling populations diverged in terms of four of six leaf anatomy traits (relative thickness of tissue layers and stomatal density), as well as leaf size and shape, specific leaf area (SLA), leaf internode distances, disparity in blade–petiole angles, canopy spread: height ratios, standing leaf numbers, summer (July) photosynthetic light curve shapes, and growth rates. Saplings showed significant sun/shade differences in fewer characters: leaf thickness, SLA, leaf overlap, disparity in bladepetiole angles, standing leaf numbers, stem volume and branching angle (first-order branches only), and summer photosynthesis. In trees, leaf anatomy was insensitive to light environment, but leaf length, width, and SLA, disparities in bladepetiole angles, and summer maximal photosynthetic rates varied among sun and shade leaf populations. Seedling and sapling photosynthetic rates were significantly depressed in winter (December), while photosynthetic rates in tree leaves did not differ in winter and summer. Seasonal and ontogenetic changes in response to light environment are apparent at several levels of biological organization in R. mangle, within constraints of its architectural baiiplan. Such variation has implications for models of stand carbon gain, and suggest that response flexibility may change with plant age.     

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     

Light,temperature and tocopherol status influence foliar vascular anatomy and leaf function in Arabidopsis thaliana

This study addressed whether the winter annual Arabidopsis thaliana can adjust foliar phloem and xylem anatomy both differentially and in parallel. In plants acclimated to hot vs cool temperature, foliar minor vein xylem‐to‐phloem ratio was greater, whereas xylem and phloem responded concomitantly to growth light intensity. Across all growth conditions, xylem anatomy correlated with transpiration rate, while phloem anatomy correlated with photosynthetic capacity for two plant lines (wild‐type Col‐0 and tocopherol‐deficient vte1 mutant) irrespective of tocopherol status. A high foliar vein density (VD) was associated with greater numbers and cross‐sectional areas of both xylem and phloem cells per vein as well as higher rates of both photosynthesis and transpiration under high vs low light intensities. Under hot vs cool temperature, high foliar VD was associated with a higher xylem‐to‐phloem ratio and greater relative rates of transpiration to photosynthesis. Tocopherol status affected development of foliar vasculature as dependent on growth environment. The most notable impact of tocopherol deficiency was seen under hot growth temperature, where the vte1 mutant exhibited greater numbers of tracheary elements (TEs) per vein, a greater ratio of TEs to sieve elements, with smaller individual sizes of TEs, and resulting similar total areas of TEs per vein and transpiration rates compared with Col‐0 wild‐type. These findings illustrate the plasticity of foliar vascular anatomy acclimation to growth environment resulting from independent adjustments of the vasculature's components.     

七种阔叶常绿植物叶片的生态解剖学研究

对 7种常绿阔叶植物叶片的解剖学特征的观察结果表明 ,它们的叶片在结构上均表现出典型的旱生特点 :异面叶 ,上表皮细胞较厚且排列紧密 ,具发达的角质膜 ,无气孔器分布 ,下表皮细胞较小 ,气孔器密度较大 ;栅栏组织细胞层数较多 ,排列紧密 ,海绵组织细胞排列极其疏松 ,并形成通气组织 ;中脉及其输导组织和机械组织发达。越冬叶和越夏叶在解剖结构上存在一定的差异 :前者在角质膜 ,叶片厚度和栅栏组织厚度等方面比后者厚 ,而在气孔密度和输导组织方面则比后者略有减少或不发达。这些差异 (发育可塑性 )是常绿阔叶植物适应冬、夏季截然不同的两种生境 (生境的时间异质性 )的表现形式     

Low temperature acclimation of photosynthetic capacity and leaf morphology in the context of phloem loading type

Carbon export from leaf mesophyll to sugar-transporting phloem occurs via either an apoplastic (across the cell membrane) or symplastic (through plasmodesmatal cell wall openings) pathway. Herbaceous apoplastic loaders generally exhibit an up-regulation of photosynthetic capacity in response to growth at lower temperature. However, acclimation of photosynthesis to temperature by symplastically loading species, whose geographic distribution is particularly strong in tropical and subtropical areas, has not been characterized. Photosynthetic and leaf anatomical acclimation to lower temperature was explored in two symplastic (Verbascum phoeniceum, Cucurbita pepo) and two apoplastic (Helianthus annuus, Spinacia oleracea) loaders, representing summer- and winter-active life histories for each loading type. Regardless of phloem loading type, the two summer-active species, C. pepo and H. annuus, exhibited neither foliar anatomical nor photosynthetic acclimation when grown under low temperature compared to moderate temperature. In contrast, and again irrespective of phloem loading type, the two winter-active mesophytes, V. phoeniceum and S. oleracea, exhibited both a greater number of palisade cell layers (and thus thicker leaves) and significantly higher maximal capacities of photosynthetic electron transport, as well as, in the case of V. phoeniceum, a greater foliar vein density in response to cool temperatures compared to growth at moderate temperature. It is therefore noteworthy that symplastic phloem loading per se does not prevent acclimation of intrinsic photosynthetic capacity to cooler growth temperatures. Given the vagaries of weather and climate, understanding the basis of plant acclimation to, and tolerance of, low temperature is critical to maintaining and increasing plant productivity for food, fuel, and fiber to meet the growing demands of a burgeoning human population.     

Effect of UV-C irradiation on stilbene synthase localization in young grape plants

Stilbene synthase (STS) is a pivotal enzyme that catalyzes the biosynthesis of resveratrol and could be induced by UV-C irradiation. However, at present the effect of UV-C irradiation on tissue and subcellular localization of STS is not studied. In this work, subcellular localization of STS in young grape (Vitis vinifera L. cv. Cabernet Sauvignon) plants exposed to UV-C was examined immunohistochemically using a polyclonal antibody raised against grape berry STS. The immunohistochemical analysis showed that the UV-induced STS occurred in palisade tissues of grape leaves and phloem tissues of grape leaf veins, stems, and roots. At the subcellular level, the enhanced STS stimulated by UV-C light was visualized in the cell walls, chloroplasts (plastids), cytoplasm, and nucleus of the phloem (stems and roots), while only in chloroplasts in mesophyll cells. This distribution pattern of STS arising in response to UV-C irradiation may be closely involved in its defense function, which needs much more in-depth evidence to confirm.     

Leaf development and phloem transport in Cucurbita pepo: Maturation of the minor veins

Summary Young leaves of Cucurbita pepo L. were examined by whole-leaf autoradiography and serial paradermal sections were examined by light microscopy to determine whether commencement of sugar export depends upon the minor vein phloem achieving structural maturity. Maturation of these veins develops progressively from the largest toward the smallest elements with the minor veins in the distal region of the leaf maturing before those in the proximal region. Commencement of sugar export is coincident with maturation of the abaxial phloem of the minor veins delimiting the areoles. The abaxial phloem elements of the larger minor veins, which are probably capable of vein loading too but border only relatively few areoles, mature before export starts. The adaxial phloem surrounding the areoles and the xylem elements, mature in advance of the abaxial phloem and well before the beginning of sugar export. It is therefore considered unlikely that structural development alone directly governs the initiation of export. The results suggest that some other rate controlling step is involved.     

Seasonal and diurnal changes in photosynthetic limitation of young sweet orange trees

This study tests the hypothesis that potted sweet orange plants show a significant variation in photosynthesis over seasonal and diurnal cycles, even in well-hydrated conditions. This hypothesis was tested by measuring diurnal variations in leaf gas exchange, chlorophyll fluorescence, leaf water potential, and the responses of CO₂ assimilation to increasing air CO₂ concentrations in 1-year-old ‘Valência’ sweet orange scions grafted onto ‘Cleopatra’ mandarin rootstocks during the winter and summer seasons in a subtropical climate. In addition, diurnal leaf gas exchange was evaluated under controlled conditions, with constant environmental conditions during both winter and summer. In relation to our hypothesis, a greater rate of photosynthesis is found during the summer compared to the winter. Reduced photosynthesis during winter was induced by cool night conditions, as the diurnal fluctuation of environmental conditions was not limiting. Low air and soil temperatures caused decreases in the stomatal conductance and in the rates of the biochemical reactions underlying photosynthesis (ribulose-1,5-bisphosphate (RuBP) carboxylation and RuBP regeneration) during the winter compared to the values obtained for those markers in the summer. Citrus photosynthesis during the summer was not impaired by biochemical or photochemical reactions, as CO₂ assimilation was only limited by stomatal conductance due to high leaf-to-air vapor pressure difference (VPD) during the afternoon. During the winter, the reduction in photosynthesis during the afternoon was caused by decreases in RuBP regeneration and stomatal conductance, which are both precipitated by low night temperature.     

Photosynthesis and water relations of well-watered orange plants as affected by winter and summer conditions

The aim of this study was to evaluate how the summer and winter conditions affect the photosynthesis and water relations of well-watered orange trees, considering the diurnal changes in leaf gas exchange, chlorophyll (Chl) fluorescence, and leaf water potential (Ψ) of potted-plants growing in a subtropical climate. The diurnal pattern of photosynthesis in young citrus trees was not significantly affected by the environmental changes when compared the summer and winter seasons. However, citrus plants showed higher photosynthetic performance in summer, when plants fixed 2.9 times more CO₂ during the diurnal period than in the winter season. Curiously, the winter conditions were more favorable to photosynthesis of citrus plants, when considering the air temperature (< 29 °C), leaf-to-air vapor pressure difference (< 2.4 kPa) and photon flux density (maximum values near light saturation) during the diurnal period. Therefore, low night temperature was the main environmental element changing the photosynthetic performance and water relations of well-watered plants during winter. Lower whole-plant hydraulic conductance, lower shoot hydration and lower stomatal conductance were noticed during winter when compared to the summer season. In winter, higher ratio between the apparent electron transport rate and leaf CO₂ assimilation was verified in afternoon, indicating reduction in electron use efficiency by photosynthesis. The high radiation loading in the summer season did not impair the citrus photochemistry, being photoprotective mechanisms active. Such mechanisms were related to increases in the heat dissipation of excessive light energy at the PSII level and to other metabolic processes consuming electrons, which impede the citrus photoinhibition under high light conditions.     

Physiological and anatomical changes induced by drought in two olive cultivars (cv Zalmati and Chemlali)

Photosynthetic gas exchange, vegetative growth, water relations and fluorescence parameters as well as leaf anatomical characteristics were investigated on young plants of two Olea europaea L. cultivars (Chemlali and Zalmati), submitted to contrasting water availability regimes. Two-year-old olive trees, grown in pots in greenhouse, were not watered for 2 months. Relative growth rate (RGR), leaf water potential (Ψ_LW) and the leaf relative water content (LWC) of the two cultivars decreased with increasing water stress. Zalmati showed higher values of RGR and LWC and lower decreased values of Ψ_LW than Chemlali, in response to water deficit, particularly during severe drought stress. Water stress also caused a marked decline on photosynthetic capacity and chlorophyll fluorescence. The net photosynthetic rate, stomatal conductance, transpiration rate, the maximal photochemical efficiency of PSII (F _v/F _m) and the intrinsic efficiency of open PSII reaction centres (F′ _v/F′ _m) decreased as drought stress developed. In addition, drought conditions, reduced leaf chlorophyll and carotenoids contents especially at severe water stress. However, Zalmati plants were the less affected when compared with Chemlali. In both cultivars, stomatal control was the major factor affecting photosynthesis under moderate drought stress. At severe drought-stress levels, the non-stomatal component of photosynthesis is inhibited and inactivation of the photosystem II occurs. Leaf anatomical parameters show that drought stress resulted in an increase of the upper epidermis and palisade mesophyll thickness as well as an increase of the stomata and trichomes density. These changes were more characteristic in cv. ‘Zalmati’. Zalmati leaves also revealed lower specific leaf area and had higher density of foliar tissue. From the behaviour of Zalmati plants, with a smaller reduction in relative growth rate, net assimilation rate and chlorophyll fluorescence parameters, and with a thicker palisade parenchyma, and a higher stomatal and trichome density, we consider this cultivar more drought-tolerant than cv. Chemlali and therefore, very promising for cultivation in arid areas.     

Photosynthetic and structural acclimation to light direction in vertical leaves of Silphium terebinthinaceum

The azimuth of vertical leaves of Silphium terebinthinaceum profoundly influenced total daily irradiance as well as the proportion of direct versus diffuse light incident on the adaxial and abaxial leaf surface. These differences caused structural and physiological adjustments in leaves that affected photosynthetic performance. Leaves with the adaxial surface facing East received equal daily integrated irradiance on each surface, and these leaves had similar photosynthetic rates when irradiated on either the adaxial or abaxial surface. The adaxial surface of East-facing leaves was also the only surface to receive more direct than diffuse irradiance and this was the only leaf side which had a clearly defined columnar palisade layer. A potential cost of constructing East-facing leaves with symmetrical photosynthetic capcity was a 25% higher specific leaf mass and increased leaf thickness in comparison to asymmetrical South-facing leaves. The adaxial surface of South-facing leaves received approximately three times more daily integrated irradiance than the abaxial surface. When measured at saturating CO₂ and irradiance, these leaves had 42% higher photosynthetic rates when irradiated on the adaxial surface than when irradiated on the abaxial surface. However, there was no difference in photosynthesis for these leaves when irradiated on either surface when measurements were made at ambient CO₂. Stomatal distribution (mean adaxial/abaxial stomatal density = 0.61) was unaffected by leaf orientation. Thus, the potential for high photosynthetic rates of adaxial palisade cells in South-facing leaves at ambient CO₂ concentrations may have been constrained by stomatal limitations to gas exchange. The distribution of soluble protein and chlorophyll within leaves suggests that palisade and spongy mesophyll cells acclimated to their local light environment. The protein/chlorophyll ratio was high in the palisade layers and decreased in the spongy mesophyll cells, presumably corresponding to the attentuation of light as it penetrates leaves. Unlike some species, the chlorophyll a/b ratio and the degree of thylakoid stacking was uniform throughout the thickness of the leaf. It appears that sun-shade acclimation among cell layers of Silphium terebinthinaceum leaves is accomplished without adjustment to the chlorophyll a/b ratio or to thylakoid membrane structure.     

A comparison of leaf physiology and anatomy of Quercus (section Erythrobalanus-Fagaceae) species in different light environments

Physiological and anatomical attributes of leaves were examined of three species of Quercus section Erythrobalanus. All three species occur in moist temperate deciduous forests of eastern North America. Seedlings of each species were grown in different light conditions for comparison. The attributes measured were net photosynthesis, stomatal conductivity, blade and cuticle thickness, stomatal density, thickness of upper and lower epidermis, and thickness of palisade mesophyll. The results generally demonstrate the close association between anatomical adaptations and efficiency of physiological processes; they also elucidate the distribution patterns of the three Quercus species across the forest topography in southern New England. The most drought-tolerant and light-demanding species, Q. velutina (Lam.), exhibited the greatest leaf anatomical plasticity, the highest net photosynthesis in the different light conditions, and the lowest stomatal area per unit area of leaf. The most drought-intolerant species, Q. rubra (L.), exhibited the least leaf anatomical plasticity, the lowest net photosynthesis in the different light conditions, and the highest stomatal area per unit area of leaf. Quercus coccinea (Muenchh.) usually exhibited values that were intermediate between Q. rubra and Q. velutina.     

陕北黄土高原优势植物叶片解剖结构的生态适应性

以陕北黄土高原地区7种优势植物为材料,比较了它们沟间地和沟谷地植株叶片解剖结构的差异,以揭示该地区优势植物对干旱强光照环境的生态适应性.结果表明:(1)7种植物各自具有抵抗黄土高原干旱强光照环境的特殊解剖结构:白羊草(Bothriochloa ischaemum)叶上表皮具有发达的泡状细胞,叶内具有花环结构;长芒草(Stipa bungeana) 叶上表皮下陷形成气孔窝,表皮下具有2至多层的厚壁组织;猪毛蒿(Artemisia scoparia)叶具有贮水组织、分泌组织和环栅型叶肉细胞;铁杆蒿(Artemisia sacrorum)叶肉全特化为栅栏组织;茭蒿(Artemisia giraldii)具有双栅型叶肉细胞和分泌结构;达乌里胡枝子(Lespedeza daurica)叶具有发达的粘液细胞;杠柳(Periploca sepium)叶表皮具厚蜡质层.(2)与沟谷地植物叶片结构相比,干旱强光照的沟间地植物叶片厚度、叶上表皮角质层厚度、栅栏组织厚度、贮水组织厚度增加,上表皮细胞体积、韧皮部面积增大,而木质部面积、木质部面积/韧皮部面积缩小.(3)叶片变异系数可反映植物适应环境的潜在能力,7种植物综合变异系数由高到低依次为猪毛蒿、铁杆蒿、茭蒿、白羊草、达乌里胡枝子、长芒草、杠柳.潜在适应能力最强的猪毛蒿已成为陕北黄土高原地区植被生态恢复的先锋物种之一.     

玉龙雪山不同海拔草血竭叶片形态与解剖结构的比较研究

采用常规石蜡制片技术和显微观察方法,对生长于滇西北玉龙雪山3 000~4 600m之间9个不同海拔高度的草血竭(Polygonum paleaceum)叶片表型及解剖结构进行比较研究,其目的是揭示该植物对不同海拔环境条件的响应特征,进而探讨植物对全球气候变化的生态适应性。结果显示:(1)草血竭叶片长度、宽度、长宽比均与海拔呈显著负相关关系,其降幅分别达84.64%、74.87%和37.50%。(2)草血竭叶片为异面叶,由表皮、叶肉、叶脉三部分组成;表皮细胞单层,角质层厚;叶肉的栅栏组织具2层或3层细胞;叶脉中的维管束呈1圈不连续的环状排列。(3)叶片厚度、栅栏组织厚度、海绵组织厚度、主脉厚度、以及主脉中维管束的长度和宽度均与海拔呈显著正相关关系,增幅分别达38.32%、47.30%、47.39%、156.46%、173.51%及337.42%;上表皮厚度、下表皮厚度、叶片紧密度、叶片疏松度、栅海比以及维管束的个数与海拔梯度之间的相关性则均不显著。研究表明,随着海拔升高,草血竭叶片形态结构的改变主要是有利于植物降低蒸腾,增强储水性和提高光合效率。     

Anatomy ofZea mays andGlycine max seedlings treated with triazole plant growth regulators

Soil drenches containing 250 μg of paclobutrazol or uniconazol (50 ml of a 17 μM solution) reduced the height of both corn (Zea mays L. cv. How.Sweet It Is) and soybean (Glycine max (L.) Merr. cv. A2) seedlings. With corn, uniconazol was considerably more active than paclobutrazol in reducing height whereas with soybean both compounds had similar dwarfing effects. The compounds increased foliar chlorophyll content and leaf thickness in soybean but had no effect on these parameters in corn. The increase in leaf thickness with soybean was due primarily to an increase in the thickness of the palisade cell layer. Chloroplast size and ultrastructure of both species were unaffected by the compounds. The growth regulators increased root diameter in both corn and soybean because of increased size of cortical parenchyma cells and particularly in soybean because of radial rather than longitudinal growth of the first few layers of the cortical parenchyma.