Ontogenetic changes in stomatal size and conductance of sunflowers

The ontogenetic changes in stomatal size, frequency and conductance (g_s) on abaxial and adaxial leaf surfaces of sunflower plants (Helianthus annuus L. Russian Mammoth) were examined under controlled environmental conditions. The stomatal frequency on the adaxial and abaxial leaf surfaces decreased with leaf ontogeny and insertion level. The ratio of adaxial to abaxial stomatal frequency did not change with leaf ontogeny and insertion level, and 42–44% of total stomata was apportioned to the adaxial surface. Ontogenetic changes in stomatal pore length were detected and increased with ontogenesis. The stomatal length of both leaf surfaces had linear relationships with leaf area. Ontogenetic changes in g_s were similar between the two surfaces. However the adaxial g_s was lower than abaxial g_s in leaves of higher insertion levels. Conductance had a linear relationship with width x frequency but not with pore area.     

Difference in light-induced increase in ploidy level and cell size between adaxial and abaxial epidermal pavement cells of Phaseolus vulgaris primary leaves

Changes in nuclear DNA content and cell size of adaxial andabaxial epidermal pavement cells were investigated using brightlight-induced leaf expansion of Phaseolus vulgaris plants. Inprimary leaves of bean plants grown under high (sunlight) ormoderate (ML; photon flux density, 163 µmol m^–2s^–1) light, most adaxial epidermal pavement cells hada nucleus with the 4C amount of DNA, whereas most abaxial pavementcells had a 2C nucleus. In contrast, plants grown under lowintensity white light (LL; 15 µmol m^–2 s^–1)for 13 d, when cell proliferation of epidermal pavement cellshad already finished, had a 2C nuclear DNA content in most adaxialpavement cells. When these LL-grown plants were transferredto ML, the increase in irradiance raised the frequency of 4Cnuclei in adaxial but not in abaxial pavement cells within 4d. On the other hand, the size of abaxial pavement cells increasedby 53% within 4 d of transfer to ML and remained unchanged thereafter,whereas adaxial pavement cells continuously enlarged for 12d. This suggests that the increase in adaxial cell size after4 d is supported by the nuclear DNA doubling. The differentresponses between adaxial and abaxial epidermal cells were notinduced by the different light intensity at both surfaces. Itwas shown that adaxial epidermal cells have a different propertythan abaxial ones. Key words: Cell enlargement, endopolyploidization, epidermal pavement cells, incident light intensity, leaf expansion, nuclear DNA content, Phaseolus vulgaris     

Leaf Pubescence, Water Relations and Chlorophyll Fluorescence in Two Subspecies of Lotus Creticus L.

The objective of the present study was to compare the plant morphology, water relations and photochemical efficiency of photosystem 2 in two wild. Mediterranean species Lotus creticus creticus and Lotus creticus cytisoides. L. creticus creticus showed higher density of trichomes and stomatal density on the adaxial leaf surface than L. creticus cytisoides, whereas L. creticus cytisoides showed higher stomatal density in abaxial surface than L. creticus creticus. These morphological traits promoted clear differences in leaf surface water retention and leaf reflectance. Leaf water potential and photochemical efficiency were lower in L. creticus cytisoides than in L. creticus creticus.     

Two sides of a leaf blade: <Emphasis Type="Italic">Blumeria graminis</Emphasis> needs chemical cues in cuticular waxes of <Emphasis Type="Italic">Lolium perenne</Emphasis> for germination and differentiation

Plant surface characteristics were repeatedly shown to play a pivotal role in plant–pathogen interactions. The abaxial leaf surface of perennial ryegrass (Lolium perenne) is extremely glossy and wettable compared to the glaucous and more hydrophobic adaxial surface. Earlier investigations have demonstrated that the abaxial leaf surface was rarely infected by powdery mildew (Blumeria graminis), even when the adaxial surface was densely colonized. This led to the assumption that components of the abaxial epicuticular leaf wax might contribute to the observed impairment of growth and development of B. graminis conidia on abaxial surfaces of L. perenne. To re-assess this hypothesis, we analyzed abundance and chemical composition of L. perenne ab- and adaxial epicuticular wax fractions. While the adaxial epicuticular waxes were dominated by primary alcohols and esters, the abaxial fraction was mainly composed of n-alkanes and aldehydes. However, the major germination and differentiation inducing compound, the C₂₆-aldehyde n-hexacosanal, was not present in the abaxial epicuticular waxes. Spiking of isolated abaxial epicuticular Lolium waxes with synthetically produced n-hexacosanal allowed reconstituting germination and differentiation rates of B. graminis in an in vitro germination assay using wax-coated glass slides. Hence, the absence of the C₂₆-aldehyde from the abaxial surface in combination with a distinctly reduced surface hydrophobicity appears to be primarily responsible for the failure of normal germling development of B. graminis on the abaxial leaf surfaces of L. perenne.     

Stomatal diffusion resistances and leaf growth during droughting of Lolium perenne plants selected for contrasting epidermal ridging

Groups of Lolium perenne plants selected for either deep or shallow adaxial epidermal ridging were grown in a 16 h day of 70 W m^-2 at 25°C, and either watered daily to 33% or allowed to dry to and then watered daily to 21% or to 16% soil moisture. During a 9 day experimental period, adaxial leaf resistances (r₁) were measured with a diffusion porometer four times daily, transpiration was estimated gravimetrically, and daily rates of leaf extension were recorded. Measurements were also made of minimum abaxial resistances, stomatal frequencies and lengths, and relative leaf water content (RLWC). At 33%, 21% or 16% soil moisture, leaf extension rates of deep ridged leaves were, respectively, slower, the same, and more rapid than those with shallow ridges. At 21% or 16% soil moisture, the adaxial r_l of deep-ridged was much lower than that of shallow-ridged leaves at all four sampling times. This difference was most marked on leaves below the youngest fully expanded, and was observed among older leaves even when plants were well watered. At low RLWC (< 85%), leaf resistance was greatest in leaves with shallow ridges. There was no significant difference between the leaf types in the calculated contributions of stomatal frequency or of morphology at any one pore opening, to r_l but deep-ridged leaves had more stomataonthe abaxial surface. Daily rate of plant water loss was directly correlated (r=+ 0.86, P < 0.01) with mean daily maximum stomatal conductance (1/r_l), and rate of leaf extension negatively with maximum r_l. It is suggested that stomata operating in the concavity formed by deep ridges open wider and are less responsive to internal changes in, for example, leaf water status, than those on shallow-ridged leaves because of a more humid microenvironment at the epidermal surface. The results are discussed in relation to the concept of ‘water-savers’ and ‘water-spenders’ and its application to breeding for dry conditions.     

西安市常见绿化植物叶片润湿性能及其影响因素

利用接触角测定仪测定了西安市21种常见绿化植物叶片表面的接触角,探讨了叶片表面特性如蜡质、绒毛、气孔对接触角的影响。结果表明,植物叶片正背面、物种间的接触角差异均显著,叶片正面和背面接触角大小在40°～140°。接触角大小与变异系数呈负相关,可能由于接触角小的润湿叶片在不同的生境和位置下,受到环境条件的影响较大而出现大的变异;接触角较大的非润湿性叶片,环境物质持留时间较短,对叶片形态和组成影响较小,因而出现小的变异。植物叶片表面的接触角随蜡质含量的升高而增大。表皮蜡质去除后大部分叶片接触角明显降低,尤其是疏水性较强的银杏(Ginkgo biloba)、月季(Ro-sa chinensis)和紫叶小檗(Berberis thunbergii)。女贞(Ligustrum lucidum)正背面、加杨(Popu-lus canadensis)背面等亲水型的叶片蜡质去除后接触角反而增大。叶片绒毛的多少及其形态、分布方式对接触角具有重要的影响,不同的作用方式表现出润湿和不润湿的特征,人为将其去除可以增加叶片的润湿性。背面气孔密度与气孔长度、保卫细胞长度呈负相关;接触角则与气孔密度呈负相关,与气孔长度呈正相关。     

西藏东达山3种嵩草属植物气孔特征沿海拔的变化

嵩草属(Kobresia)植物是藏东南高山草甸的优势种和建群种,对该区畜牧业发展和维持生态系统平衡起着重要作用。选择西藏左贡县东达山为研究地点,从林线开始,海拔每升高约100m设置1个样带直至高山草甸分布边缘,共8个样带,调查各样带中物种的组成及盖度,并依据相对盖度和相对频度计算3种嵩草植物矮生嵩草(K.humilis)、线叶嵩草(K.capillifolia)和大花嵩草(K.macrantha)在群落中的重要值,同时取样观察它们叶片远、近轴面表皮细胞形态,测量气孔长度及保卫细胞宽度,计算气孔密度,探讨嵩草属植物对海拔梯度的适应性。结果表明:(1)3种嵩草属植物叶表皮细胞均呈波浪状,气孔器仅分布于远轴面,近轴面无气孔器分布。(2)3种嵩草属植物气孔密度沿海拔梯度的变化均呈单峰曲线分布格局,且在海拔4 537m样带处达到最大值,并表现为矮生嵩草(777.6个/mm2)线叶嵩草(476.4个/mm2)大花嵩草(414.3个/mm2)。(3)随海拔的增加,矮生嵩草和线叶嵩草气孔长度显著增大(P0.05),而保卫细胞宽度显著减小;但大花嵩草气孔长度随海拔的升高而显著减小,保卫细胞宽度基本保持不变。(4)矮生嵩草和线叶嵩草气孔密度、长度和保卫细胞宽度与海拔梯度均显著相关,气孔特征对海拔梯度变化的敏感程度高,与其在群落中重要值高的分布特征一致;而大花嵩草仅气孔密度和长度与海拔梯度显著相关,气孔特征对海拔梯度变化的敏感性低,与其在群落中重要值低的分布特征一致;嵩草属植物气孔密度、长度和保护细胞宽度与海拔梯度之间的相关性,反映出它们在海拔梯度上对生境的适应程度。可见,3种嵩草属植物气孔特征对海拔梯度上生境变化的适应性不同,从而影响它们在群落中的分布范围和物种优势度,其中矮生嵩草和线叶嵩草对环境变化敏感,而大花嵩草对环境变化相对不敏感;保卫细胞宽度与气孔长度同样对植物适应环境变化起重要作用。     

Photosynthetic energy storage in aquatic leaves measured by photothermal deflection

In a study of photosynthetic energy storage efficiency (ES), the adaxial surface of the leaves of Vallisneria americana exhibited the highest ES values (22%) of the four aquatic plants examined. V. americana leaves have a dispersed structure and it was possible to measure the energy storage properties of the epidermal cells independently of the rest of the leaf. The abaxial epidermis had a higher value of ES at zero light fluence than the adaxial epidermis but ES in the abaxial epidermis declined much more rapidly with light fluence. Thus the abaxial epidermis is more suited to lower light fluences than the adaxial epidermis. ES declined as the pH rose from 4.0 to 8.0 at a constant dissolved inorganic carbon concentration. This paralleled the change from carbon dioxide to bicarbonate and suggests that these leaves utilise CO₂ more efficiently than bicarbonate. ES increased by about 50% at pH 8.0 as leaf sections further from the leaf tip were examined which demonstrates that the older epidermal cells are less well able to use bicarbonate. Exposure to 30 min of a saturating light fluence caused the epidermal chloroplasts to move from the periclinal walls to the anticlinal walls. This decreased the photothermal signal by increasing the thermal diffusion distance and lowering the light fluence due to greater chloroplast shading. The latter effect increased ES. It appears that chloroplast movement could assist the epidermis to survive harmful light fluences.Abbreviations APW artificial pond water; atrazine- (2-chloro-4-ethylamino-6-(isopropylamine)-s-triazine) - DCMU (3-(3,4 dichlorophenyl)-1,1-dimethyl urea) - DIC dissolved inorganic carbon - DMSO dimethyl sulfoxide - ES energy storage efficiency - HEPES N-[2-Hydroxyethyl]piperazine-N-[2-ethanesulfonic acid] - MOPS 3-[N-Morpholino]propanesulfonic acid - PD photothermal deflection     

Diurnal variation in leaf water potential,stomatal conductance,and irradiance of winter crop under different moisture levels

Mustard (Brassica juncea) Coss., chickpea (Cicer arietinum L. and barley (Hordeum vulgare) L. were grown under different moisture levels. Diurnal changes in leaf water potential showed lower values and higher fluctuation in mustard, chickpea and barley grown with no irrigation as compared to one supplemental irrigation. Diurnal maximum of adaxial stomatal conductance in mustard and barley was higher under one irrigation treatment. In mustard stomatal conductance of abaxial surface of leaf remained higher than adaxial surface of leaf throughout the day, whereas the reverse was true in barley. Also the leaf and soil temperature and reflectance were slightly higher in all the three crops under no irrigation.     

Abaxial and Adaxial Stomatal Density, Stomatal Conductances and Water Status of Bean Primary Leaves as Affected by Paclobutrazol

The plant growth retardant, paclobutrazol at 8.5 or 17.0 μM concentrations effectively inhibited the stem elongation and primary leaf expansion of bean seedlings. Although the retardant reduced the relative water content in well-watered plants, the water and pressure potentials remained high in the primary leaves. K⁺, Na⁺, Mg²⁺ and Ca²⁺ contents in the primary leaves of the paclobutrazol-treated plants were not significantly different from those in the control. The stomatal density increased on both surfaces but the length of guard cells was not reduced significantly on the adaxial epidermes of the paclobutrazol-treated primary leaves. The inhibitory effect of paclobutrazol on the abaxial stomatal conductances became more pronounced with time during the light period but the adaxial surfaces displayed similar or slightly higher conductances than those of the control. The transpiration rate on a unit area basis did not change significantly or increased in the treated leaves thus the reduced water loss of paclobutrazol-treated plants was due to the reduced leaf area. Stomatal conductances of the adaxial surfaces responded more intensively to exogenous abscisic acid and the total leaf conductance decreased faster with increasing ABA concentration in the control than in the paclobutrazol-treated leaves. Paclobutrazol, an effective inhibitor of phytosterol biosynthesis, not only amplified the stomatal differentiation but increased the differences between the adaxial and abaxial stomatal conductances of the primary leaves.     

Leaf growth, stomatal diffusion resistances and photosynthesis during droughting of Lolium perenne populations selected for contrasting stomatal length and frequency

Lolium perenne selection lines with high calculated stomatal resistances to diffusion (r_s) as a result of either few or short stomata, maintained leaf extension rates and photosynthetic rates longer than selection lines with low resistances when deprived of water. There were no significant differences between high and low r_s plants in light saturated CO₂ uptake of turgid attached leaves. When grown in soil drying to 21% moisture, plants with low calculated r_s exhibited lower minimum leaf resistances (r_l) than those with high, measured with a diffusion porometer, on all except the last day. The daily maximum r_l (1.5 h after the start of the light period) became greater among low than high r_s plants as the difference in rate of leaf extension between the two groups of plants increased with drying soil. Rate of leaf extension was negatively correlated with daily maximum r_l and started to decline when relative leaf water content (RLWC), at 5 h after the start of the light period, fell below about 88%. Transpiration rate of plants grown in different soil moisture regimes was correlated (r=+ 0.83, P < 0.01) with mean maximum adaxial leaf conductance (reciprocal of resistance). There was a highly significant correlation (r=+ 0.62, P < 0.01) between calculated adaxial r_s and mean minimum measured r_l among plants growing at high or intermediate soil moisture, but not at low. Therefore, some random variation in minimum r_l, even with adequate moisture, seemed to be unaccounted for by variation in stomatal numbers or size. Selection for increased numbers of adaxial stomata also resulted in more on the abaxial surface, but mean adaxial/abaxial ratio in the ‘frequent’ stomata plants was still only about 9:1.     

DEVELOPMENTAL CHANGES IN THE COTYLEDONS OF LUPINUS LUTEUS L. DURING AND AFTER GERMINATION

Cotyledons of Lupinus luteus were sampled from 1 to 21 days after sowing and processed for light microscopy and sodium dodecyl sulphate polyacrylamide gel electrophoresis (SDS-PAGE). Length, width, and surface area of the cotyledons increased gradually until day 10. The thickness of the cotyledons increased from day 7 to day 12 and decreased thereafter. Morphometric analyses showed that the increase in length, width, and thickness of the cotyledon was due to cell expansion, and the decrease in thickness of the cotyledon was due to the decrease in the length of abaxial cells and in the total number of cells. Mesophyll development accompanied schizogenous and lysigenous air space formation. There were two structurally distinct types of protein bodies. Protein bodies in five to six layers of cells on the abaxial side did not contain globoids, while globoids were prominent in protein bodies in the center and adaxial side. Storage protein mobilization occurred first in the abaxial side of the cotyledon and proceeded toward the adaxial side. SDS-PAGE analysis showed that proteins ranged from 97 to 14 kD. High molecular weight α- and ß-conglutinins were more abundant in the abaxial region, whereas γ-conglutinin occurred in both abaxial and adaxial regions. In addition, there were five minor bands between 97 and 43 kD unique to abaxial region and five minor bands between 43 and 14 kD unique to adaxial region in the nonreduced protein profiles. The α- and ß-conglutinins began to decrease after imbibition and disappeared by day 7 after sowing. At this stage the subunits of ribulose-l,5-bisphosphate carboxylase/oxygenase and four new minor bands appeared.     

The relationship between water availability and anatomical characters in Carex hirta

The effects of edaphic moisture in anatomical characters were evaluated in two different populations of Carex hirta L. with three watering treatment for 6 months to evaluate stability, and determined taxonomic value. Water availability increased (p < 0.001) leaf thickness from 239 to 289 μm, metaxylem vessel diameter from 17 to 23 μm, air cavity size from 10 to 24% and adaxial epidermal cell height from 18 to 34 μm, and abaxial from 11 to 16 μm, adaxial epidermal cell length from 54 to 105 μm, and abaxial from 35 to 86 μm, and adaxial epidermal cell width from 20 to 33 μm, and abaxial from 15 to 23 μm. Stomatal index and the number of cells in the girder of sclerenchyma did not vary with water availability, hence these traits have taxonomic value. Other characters (the length and amplitude of wall undulations in the epidermal cells, the number of bulliform cells) have a doubtful relation with water availability, because they are variable even in constant homogeneous conditions.     

中国卫矛属植物叶表皮微形态特征及其分类学意义

利用光学显微镜和扫描电子显微镜对中国卫矛属(Euonymus L.)24种以及沟瓣属（Glyptopetalum Thw.）3种植物的叶表皮微形态特征进行了观察研究,测量并统计了气孔器长宽比、密度和气孔指数。结果显示,两属植物叶表皮微形态特征基本一致：叶上、下表皮细胞形状多为多边形,垂周壁平直-弓形;气孔器通常仅分布在叶下表皮;均具有无规则型气孔器,部分种还具环列型气孔器。扫描电镜下角质膜纹饰、表皮细胞界限及气孔特征在一些种间存在差异,可以作为区分部分近缘种类的参考依据。     

Effects of experimental warming on stomatal traits in leaves of maize (Zea may L.)

We examined the warming effects on the stomatal frequency, stomatal aperture size and shape, and their spatial distribution pattern of maize (Zea may L.) leaves using a light microscope, an electron scanning microscope, and geostatistic techniques. A field manipulative experiment was conducted to elevate canopy temperature by 2.08°C, on average. We found that experimental warming had little effect on stomatal density, but significantly increased stomatal index due to the reduction in the number of epidermal cells under the warming treatment. Warming also significantly decreased stomatal aperture length and increased stomatal aperture width. As a result, warming significantly increased the average stomatal aperture area and stomatal aperture circumference. In addition, warming dramatically changed the stomatal spatial distribution pattern with a substantial increase in the average nearest neighbor distance between stomata on both adaxial and abaxial surfaces. The spatial distribution pattern of stomata was scale dependent with regular patterns at small scales and random patterns at larger scales on both leaf surfaces. Warming caused the stomatal distribution to become more regular on both leaf surfaces with smaller L(t) values (Ripley's K‐function, L(t) is an expectation of zero for any value of t) in the warming plots than the control plots.     

Variation in Selection for. Leaf Water Conductance in Relation to Growth and Stomatal Dimensions in Lolium perenne L.

A simple, rapid technique for direct selection for leaf waterconductance (LWC) in two populations of Lolium perenne L. isdescribed. Measurements were made with a diffusion porometerin growth rooms on the youngest fully expanded leaf of eachtiller. Considerable variation in LWC was found between 100genotypes of each population. Most of the variation in totalLWC was attributed to variation in adaxial LWC and it was shownthat ignoring abaxial LWC gave very similar ranking of genotypesto those using total LWC. Selection criteria were then establishedwhich maximised the repeatability of LWC measurements and allowedselection of groups of plants significantly different from oneanother in LWC. The variation in LWC was not related eitherto stomatal length or number, or to growth rate. Lolium perenne L., perennial ryegrass, abaxial and adaxial, leaf water conductance, selection, diurnal rhythms, ontogenetic changes, growth, stomatal dimensions     

Leaf epidermal micromorphology of Cercis (Fabaceae: Caesalpinioideae)

The micromorphology of leaf epidermal cells and stomata of all eight species and one form (11 samples) of Cercis were observed by scanning electron microscopy and light microscopy. Both the adaxial and abaxial epidermal cells are polygonal or irregular in shape; the anticlinal walls are straight and arched or undulate. Two types of stomata, which occur only on the abaxial surface of the leaves, are found in the genus. The atypical paracytic type is present in only one species, Cercis chingii, and the anomocytic type is present in all other species. Interspecific differences are minor in the genus with regard to leaf epidermal characters, except for C. chingii, which is characterized by atypical paracytic‐type stomata, a two‐lipped outer stomatal rim, the highest stomatal density and undulate and densely pitted anticlinal walls in the adaxial epidermis. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 539–547.     

Effects of low level O3 exposure on leaf diffusive conductance and water-use efficiency in hybrid poplar

Abstract Young, amphistomatous hybrid poplar (Populus deltoides x trichocarpa) plants were exposed daily to either background (0.025 cm³ m^-3) or elevated (0.125 cm³ m^-3) concentrations of O₃. Levels of abaxial and adaxial leaf conductance were affected interactively by pollutant treatment, leaf age, and photon fluence rate. Consequently, conductance in O₃-treated leaves was sometimes higher and sometimes lower than in comparable control leaves, depending on leaf age or level of photon fluence rate. For example, at low photon fluence rate or in the dark, conductance was greater in O₃-treated than in control plants, while at high photon fluence rate that relationship was reversed. Exposure to O₃ also reduced the water-use efficiency and range of leaf conductance of individual leaves, and altered the relationship between the conductances of the two leaf surfaces (the ratio of abaxial to adaxial leaf conductance was increased). Furthermore, O₃ treatment resulted in diminished stomatal control of water loss; excised O₃-treated leaves had higher conductances and wilted sooner than excised control leaves of identical ages. Overall, the data indicate that exposure to O₃ resulted in impaired stomatal function.     

Stomatal and photosynthetic responses ofCichorium intybus leaves to sulfur dioxide treatment at different stages of plant development

Fifty-day-oldCichorium intybus Linn, plants were exposed to 1 ppm sulfur dioxide gas, 2 h per day for 7 consecutive days. Their leaves as well as those from the control plants were sampled at pre-flowering, flowering, and post-flowering stages to study their morphological, physiological, and biochemical responses to SO₂ stress. The number, dimensions, area, and biomass of leaves were less in the treated plants. Length and width of stomatal apertures on both epidermises were greater for leaves exposed to SO₂. The Stomata were longer on the adaxial epidermis, but shorter on the abaxial epidermis, except at the pre-flowering stage. Stomatal widths varied widely. Compared with the controls, the abaxial epidermis on treated leaves showed consistently lower stomatal densities as well as stomatal indices. This was also true for the adaxial epidermis during the post-flowering stage. The photosynthetic rate and stomatal conductance were reduced in the SO₂-exposed plants, but intercellular CO₂ concentrations increased at the pre-flowering stage and, subsequently, declined. Chlorophyll a, carotenoid, and total chlorophyll contents increased at the pre-flowering stage, and then decreased. The level of chlorophyllb was reduced throughout plant development compared with the untreated controls.