植物叶的波动性不对称对空气污染的响应及其指示功能

植物发育的波动性不对称(fluctuating asymmetry,FA)与环境之间具有密切的关系。该研究以城市常用绿化树种大叶女贞(Ligustrum lucidum Ait.)和小叶女贞(Ligustrum quihoui Carr.)为对象,在西安市区9个空气质量监测点采集了两树种成熟叶片样品,以叶脉为对称轴测定叶左右两边的半宽度、半周长和半面积,计算其相应FA,考察植物叶片FA对空气环境条件变化的响应特征,探讨叶FA作为空气质量指标的可行性。结果表明:大叶女贞叶周长(FAP)、宽度(FAW)和面积(FAA)波动性不对称分别为0.051、0.063和0.082,小叶女贞分别为0.043、0.082和0.102,均以FAA最大,FAP最小;FAP、FAW和FAA在树种间、立地环境间均具有极显著的差异;两树种的FAP与空气中SO_2浓度之间存在显著相关性。可见,大叶女贞和小叶女贞叶的波动性不对称特征对不同环境条件具有敏感的反应,具有作为生物监测指标的潜力,但在合适监测物种和叶特征指标的选择、样本数确定以及FA与空气特征污染物的关系等方面有待深入探讨。     

青藏高原和内蒙古高原典型草地植物叶片肾型和哑铃型气孔器气孔特征及其与环境的关系

该研究利用植物制片技术,以中国青藏高原和内蒙古高原典型草地常见种或优势种植物的叶片为研究对象,通过比较分析叶片哑铃型气孔器和肾型气孔器的特征及其与环境因子的关系,揭示植物叶片两类气孔器对环境因子的响应策略。结果表明:(1)叶片哑铃型气孔器气孔指标的变异系数小于肾型气孔器。(2)叶片哑铃型气孔器气孔指标与环境气候指标的关系弱于肾型气孔器。(3)叶片哑铃型气孔器气孔特征与环境关系在叶片上下表面之间存在显著差异,而肾型气孔器气孔特征与环境因子的关系在叶片上下表面之间无显著差异。(4)叶片哑铃型气孔器的气孔特征与降水关系密切,而肾型气孔器气孔特征与温度关系密切。(5)同一种气孔器的气孔特征在两个地区(青藏高原和内蒙古高原)间存在显著差异。研究认为,肾型气孔器和哑铃型气孔器的气孔特征及其与环境之间的关系存在差异,在分析气孔特征时有必要将肾形与哑铃形保卫细胞形成的气孔器加以区分,该研究结果有助于进一步理解中国草地植物叶气孔特征对气候变化的响应与适应策略。     

赤皮青冈幼苗叶片解剖结构对干旱胁迫的响应

以浙江庆元、湖南洞口和湖南靖州3个种源的赤皮青冈1年生幼苗为材料,采用盆栽称量法控制土壤水分含量,设置土壤相对含水量分别为75%～80%(对照)、55%～60%(轻度干旱胁迫)、45%～50%(中度干旱胁迫)和30%～35%(重度干旱胁迫),研究不同程度干旱胁迫对赤皮青冈幼苗叶片解剖结构和蒸腾作用的影响.结果表明: 干旱胁迫下,3个种源赤皮青冈幼苗叶片的总厚度、上下表皮厚度和栅栏组织厚度均显著降低.栅栏组织厚度与海绵组织厚度比(栅海比)、气孔长度和宽度随干旱程度的增强而显著减小,而气孔密度随干旱胁迫程度的增强而显著增加.干旱处理对叶片主脉厚度无显著影响,木质部厚度的变化因种源而异.叶片结构的变化引起生理功能改变,随着胁迫程度的加强,赤皮青冈幼苗的蒸腾速率显著降低.对照和各干旱处理湖南洞口种源幼苗叶片的下表皮厚度、栅海比、气孔密度显著大于其他2个种源,蒸腾速率显著小于其他2个种源,表明湖南洞口赤皮青冈对干旱胁迫的适应性较强.     

太白山栎属树种气孔特征沿海拔梯度的变化规律

气孔是植物与外界环境进行水分和气体交换的主要通道,调节植物碳同化和水分散失的平衡关系,在一定程度上反映植物对外界环境变化的适应。沿太白山北坡1100—2300 m海拔,测定4种栎属树种的气孔性状,分析气孔性状沿海拔的变化规律和其对环境因子的响应。结果表明:(1)气孔密度与气孔长度间的负相关在4个树种间均显著存在(P0.05);除栓皮栎(Quercus variabilis)外,气孔密度与潜在气孔导度指数的正相关关系均达显著水平;而气孔宽度与气孔长度之间只在栓皮栎和锐齿栎(Q. aliena var. acuteserrata)达到显著水平。(2)栓皮栎和槲栎(Q. aliena)的气孔长度和宽度随海拔升高而下降,气孔密度、潜在气孔导度指数增加,辽东栎(Q. wutaishansea)变化形式则相反;锐齿栎气孔宽度减小,其余性状沿海拔呈单峰变化,在约1600 m处气孔长度达到最小值,气孔密度和潜在气孔导度指数达到最大值。(3)与土壤因子相比,气孔性状主要受气候因素的影响。潜在气孔导度指数与大气温度、空气湿度成极显著正相关(P0.01),与降水量显著负相关(P0.05)。其中,空气相对湿度是影响潜在气孔导度指数的主要因素,能够解释气孔变异的22.9%。本研究结果对于深入认识秦岭太白山地区栎属树种对环境变化的响应和适应提供理论证据。     

NaCl胁迫对两种番茄气孔特征、气体交换参数和生物量的影响

为探讨番茄叶片气孔特征、气体交换参数和生物量对盐胁迫的响应机理,以赛棚和阿拉姆番茄为试材,通过向水培营养液中添加NaCl(0.1 mol·L^-1),在人工气候箱条件下进行为期90 d的NaCl盐胁迫处理.结果表明: NaCl胁迫导致赛棚番茄叶片的气孔密度、气孔宽度、气孔面积和气孔面积指数显著降低,降幅分别为32%、45%、25%、49%,但没有改变阿拉姆番茄叶片的气孔特征参数.同时,NaCl胁迫还导致赛棚和阿拉姆叶片气孔规则分布的空间尺度分别减少30%和43%,且赛棚品种的单个气孔最小邻域距离在盐胁迫下增加20%.另外,赛棚和阿拉姆叶片的净光合速率(P_n)、气孔导度(g_s)和蒸腾速率(T_r)在盐胁迫下均显著下降,通过气孔限制值分析发现,盐胁迫导致赛棚番茄叶片光合速率下降主要是由气孔限制因素引起的,而阿拉姆叶片则以非气孔限制因素为主导作用.盐胁迫还导致赛棚和阿拉姆番茄生物量显著降低,且地下生物量的下降幅度大于地上生物量.综合分析表明,阿拉姆的抗盐能力高于赛棚.     

中间锦鸡儿叶表皮微形态的经度格局及其影响因素

环境因子是影响植物种内变异的主要因素,而叶片是植物体最易受环境影响的器官。为了探究中间锦鸡儿植物叶表皮微形态是否具有经度变异及其影响因素,该研究在内蒙古高原调查了8个中间锦鸡儿种群叶片的气孔密度、气孔指数、表皮毛密度、气孔器长、气孔器宽,并利用回归分析探究其变异程度以及影响因子。结果表明：(1)锦鸡儿上下表皮均被有单细胞非腺毛,角质层粗糙,气孔下陷,为抗旱特征;中间锦鸡儿上表皮气孔密度、气孔指数、表皮毛密度、气孔器长和气孔器宽的均值分别为302.81个/mm²、9.94、204.81个/mm²、11.78 μm和5.95 μm,而叶片下表皮分别为272.83个/mm²、10.43、187.45 个/mm²、12.49 μm和6.18 μm。(2)中间锦鸡儿叶片的气孔密度与气孔指数随经度的增加而增加,而表皮毛密度随经度的增加却减小。(3)叶片上下表皮的可塑性指数排序均为：表皮毛密度>气孔密度>气孔指数>气孔器宽>气孔器长;中间锦鸡儿的叶表皮主要通过调节表皮毛密度及气孔密度来响应环境的变化,环境因子中日照时数、有效积温、相对湿度、土壤全碳、全磷、全氮含量显著影响叶片表皮的微形态特征。     

润楠属植物叶片和枝条水力特征的协同关系

以润楠属(Machilus) 7种植物成年个体为材料,对其进行生理指标测定,并对它们的叶片水分供需关系以及木质部纹孔特征和导水效率之间的关联进行分析。结果显示,润楠属7种植物相比原始被子植物具有更高的叶脉密度(VD),叶脉密度为9.8～14.1 mm/mm~2;气孔密度(SD)与叶脉密度呈显著正相关,说明叶片水分供需存在协同关系;气孔密度与气孔大小(GLC)呈负相关;气孔越大的叶片其膨压丧失点(TLP)的绝对值越低。枝条边材比导率(Ks)较低,为0.13～1.87 kg·m~(-1)·s~(-1)·MPa~(-1),且种间差异较大。叶脉和气孔密度均与边材比导率呈正相关。边材比导率与纹孔膜面积、纹孔口面积以及纹孔口长短轴比例相关性不显著。研究结果表明润楠属植物虽然叶脉密度较高,且木质部水分供应和叶片结构具有协同关系,但木质部解剖结构较为原始,导管多具梯形穿孔板,导水效率低,只能适应比较湿润的生境。     

大气中CO2、O3浓度升高对银杏成年叶片气孔数量特征的影响

利用开放式气室(OTC)持续观测了2个生长季(2005-2006年).在每年9月30日停止供气后,采样1次.观测高浓度CO2和O3处理的银杏(Ginkgo biloba)成年叶片气孔数量的变化规律.结果表明:在700 μmol·mol-1CO2处理条件下,叶片气孔的长度、宽度、周长和面积均明显高于对照(P<0.05);在80 μmol·mol-1O3处理条件下,气孔的长度、宽度、周长和面积却显著低于对照(P<0.01);成年叶片气孔在高浓度CO2影响下,气孔密度略有升高(P>0.05),而气孔指数显著减少(P<0.05);大气O3浓度升高对气孔密度影响不大(P>0.05),气孔指数显著减少(P<0.05);银杏成年叶片气孔的长度、宽度与气孔比密度存在一定的负相关关系,可用一元二次曲线方程较好地描述.     

土壤水分胁迫对设施番茄叶片气孔特性的影响

为了研究土壤水分胁迫对设施番茄叶片气孔特性的影响,本研究以"金粉2号"为试材,于2013年5—8月设计4个土壤水分梯度试验[正常灌溉(田间持水量的70%~80%)、轻度胁迫(田间持水量的60%~70%)、中度胁迫(田间持水量的50%~60%)和重度胁迫(田间持水量的30%~40%)],利用数码显微成像系统和数码测距软件测定了叶片气孔参数。结果表明:番茄叶片气孔长度、宽度、气孔开张度和开张比均随水分胁迫程度的增加而减小,且随着胁迫时间的延长,其减小幅度不断增大;番茄气孔长宽的减小具有同步性,但气孔长度下降更为显著;随着水分胁迫程度的增加,气孔逐渐由长卵形转变为近圆形;随着土壤水分胁迫强度的增加,气孔上表皮气孔密度逐渐增大,而气孔下表皮密度呈先减小后增大的趋势。该研究揭示了土壤水分胁迫对设施番茄叶片气孔发展的影响规律,可为设施番茄水分管理提供科学依据。     

青藏高原草地双子叶植物叶片的气孔特征研究

利用光学显微镜对青藏高原29种草地双子叶植物叶片的气孔形态与数量特征进行观察及差异显著性分析,为揭示青藏高原草地双子叶植物对高原环境的适应机制及探索气孔作为辅助分类的依据奠定理论基础。结果表明:(1)青藏高原草地双子叶植物大多数种类在叶片上、下表皮均分布有气孔,气孔随机排列,气孔器多为无规则型。(2)气孔长度(SL)较小,上、下表皮的气孔平均长度分别为26.20μm与25.56μm,且气孔密度(SD)与气孔指数(SI)相对较大。(3)不同科、属、种间叶片上、下表皮的SL、SD、SI差异均极显著。(4)叶片上、下表皮的6个气孔数量特征之间具有显著相关关系。(5)上表皮的SL、SD与不同科、属、种间显著相关,下表皮除SI与物种间相关不显著外,其他指标与科、属、种间显著相关。研究认为,青藏高原草地双子叶植物独特的气孔形态与数量特征是对高寒极端环境长期适应的结果,且气孔数量特征对植物辅助分类具有重要价值。     

Leaf developmental stability of Platanus acerifolia under urban environmental stress and its implication as an environmental indicator

The developmental stability indices,leaf width based fluctuating asymmetry (FA1),and lateral vein length based directional asymmetry (RDA1) of Platanus acerifolia were studied.All the leaves were sampled from 14 sites that were categorized based on different urban environmental stress levels (UESL) in Shanghai metropolitan,China.Besides,foliar stomatal density and stomatal length were also studied as the subsidiary indices to test the availability of developmental stability indices as the indicator under a stressful environment.Results showed seasonal variation of FA1 and RDA1 existed among the 14 sites,but the data showed significant negative correlation between FA1 and UESL (FA1=0.029-0.000 9UESL+0.000 3UESL2,r=0.766 5,P=0.001 4).However,a similar trend was not found between RDA1 and UESL.Furthermore,the significant correlation among FA1 and leaf stomatal length and stomatal density implied they could be used as indicators of urban stress levels on a small scale.It seemed that RDA1 was possibly a normal parameter during leaf development but it was unavailable for use as an indicator of urban stresses.     

Leaf developmental stability of Platanus acerifolia under urban environmental stress and its implication as an environmental indicator

The developmental stability indices, leaf width based fluctuating asymmetry (FA1), and lateral vein length based directional asymmetry (RDA1) of Platanus acerifolia were studied. All the leaves were sampled from 14 sites that were categorized based on different urban environmental stress levels (UESL) in Shanghai metropolitan, China. Besides, foliar stomatal density and stomatal length were also studied as the subsidiary indices to test the availability of developmental stability indices as the indicator under a stressful environment. Results showed seasonal variation of FA1 and RDA1 existed among the 14 sites, but the data showed significant negative correlation between FA1 and UESL (FA1 = 0.029 − 0.000 9UESL + 0.000 3UESL², r = 0.766 5, P = 0.001 4). However, a similar trend was not found between RDA1 and UESL. Furthermore, the significant correlation among FA1 and leaf stomatal length and stomatal density implied they could be used as indicators of urban stress levels on a small scale. It seemed that RDA1 was possibly a normal parameter during leaf development but it was unavailable for use as an indicator of urban stresses. __________ Translated from Chinese Journal of Ecology, 2005, 24(7): 719–723 [译自: 生态学杂志, 2005, 24(7): 719–723]     

Fluctuating asymmetry of leaves is a poor indicator of environmental stress and genetic stress by inbreeding in Silene vulgaris

Fluctuating asymmetry (FA) is often used as a measure of developmental instability and has been proposed as an indicator of both environmental and genetic stress. However, the empirical evidence for the effects of stress on FA in plants is inconsistent, and there are few controlled experimental studies. We analyzed different distance- and shape-based measures of the fluctuating asymmetry of the leaves of clonally replicated self- and cross-pollinated lineages of Silene vulgaris (Moench) Garcke (Caryophyllaceae) grown under a control and seven different stress treatments (drought, copper, simulated herbivory, and two levels of nutrient deficiency and of shade). Overall, FA differed among stress treatments, but was not generally higher under stress and even reduced in some treatments. Different measures of FA were only weakly correlated. Inbreeding increased only one measure of leaf FA, and only under high stress intensities. Our findings suggest that in S. vulgaris leaf fluctuating asymmetry does not serve as an indicator of environmental stress or of genetic stress by inbreeding.     

A modified version of fluctuating asymmetry, potential for the analysis of Aesculus hippocastanum L. compound leaves

My research interest was to create a new, simple and tractable mathematical framework for analyzing fluctuating asymmetry (FA) in Aesculus hippocastanum L. palmately compound leaves (each compound leaf with 7 obviate, serrate leaflets). FA, being random differences in the development of both sides of a bilaterally symmetrical character, has been proposed as an indicator of environmental and genetic stress. In the present paper the well-established Palmer's procedure for FA has been modified to improve the suitability of the chosen index (FA1) to be used in compound leaf asymmetry analysis. The processing steps are described in detail, allowing us to apply these modifications for the other Palmer's indices of FA as well as for the compound leaves of other plant species.     

Fluctuating asymmetry in a secondary sexual trait: no associations with individual fitness, environmental stress or inbreeding, and no heritability

It has been suggested that fluctuating asymmetry (FA) in secondary sexual traits may be a useful indicator of either individual quality or environmental stress. We tested this concept using a series of analyses of FA in male antler size in a wild red deer (Cervus elaphus) population, using four measures of size repeated across successive years on the same individuals. We found no consistent evidence of correlations between traits in levels of FA, nor of any associations between known environmental or developmental conditions. None of the four measures of FA showed a significant heritability (average h2 = 0.041), nor was there any evidence of inbreeding depression. For three of the four traits, fluctuating asymmetry did not predict either annual or lifetime breeding success. However there were significant associations between breeding success and FA in antler length. Given the series of null results in our other tests, it seems likely that this was a direct mechanistic effect rather than because measures of FA were indicative of individual quality or condition.     

Similar responses of insect herbivores to leaf fluctuating asymmetry

Fluctuating asymmetry (FA) represents small, random variation from symmetry and it has been used as an indicator of plant quality and susceptibility to herbivory. In this study, the effects of FA on the responses of distinct herbivore species belonging to several guilds were examined along an environmental gradient in south Florida. This approach was chosen because it relies on a multi-species approach to the study of fluctuating asymmetry and patterns of herbivory between and within plants along an environmental gradient of salinity and plant stress. To examine differences in FA between and within plant communities, seven plant species were investigated. Four of these plants were coastal species and three species occurred in upland communities. Levels of FA were assessed before herbivory and plants were followed for the whole herbivory season in 2006. Coastal plants exhibited significantly higher salt concentration, higher percentage of asymmetric leaves and higher asymmetry levels than upland plants. Herbivore abundance varied widely amongst the seven species studied, but quantitative syntheses of our results indicated significant and positive responses of insect herbivores to leaf asymmetry: insects were 25.11% more abundant on more asymmetric plants and stronger effects of asymmetry were observed for leaf miners compared to gall-formers. As demonstrated by other recent studies, FA might be used as a reliable stress indicator, leading to similar responses of insect herbivores to variation in leaf symmetry.     

Lizards from Urban Areas Are More Asymmetric: Using Fluctuating Asymmetry to Evaluate Environmental Disturbance

The increase in human activities that leads to wildlife decline and species extinction poses an urgent need for simple indicators of environmental stress in animal populations. Several studies have suggested that fluctuating asymmetry (FA) can be an easy, direct measure of developmental instability because it is associated to environmental stress and, as such, it can be a useful indicator of population disturbance. We examined three different morphological traits in urban and rural populations of the common wall lizard (Podarcis muralis) to test whether anthropogenic disturbance causes an increase in FA. Compared to rural populations, urban ones showed higher levels of FA in all analyzed traits, thus providing evidence that FA can respond to anthropogenic disturbance. However, we also found significant differences in FA among traits, where femoral pores and subdigital lamellae, traits with a functional relevance, were more stable developmentally compared to supracilliar granules which have no evident function. Unsigned FA [abs(right-left)] exhibited significant, but weak, positive correlations among traits, indicating that developmental noise does not have a uniform effect across characters and thus questioning the view of developmental stability as an organism-wide property. The degree of signed FA (right-left) was more similar between structurally associated traits, possibly as an outcome of morphological integration. In conclusion, our results demonstrate that FA can be a reliable indicator of disturbance provided that it is analyzed on multiple traits simultaneously and examined at the population level.     

Stability in development and variability in morphological signs in a natural population of Drosophila melanogaster: seasonal dynamics in 1999

Seasonal dynamics of developmental stability and variability of morphological traits was examined in a natural population of Drosophila melanogaster in order to compare these two parameters as indicators of temperature stress. Morphometric (thorax length and wing length) and meristic (number of sternopelural and orbital bristles) were studied. Variability was measured as the coefficient of variation. Stability of development was estimated as fluctuating asymmetry (FA) of bilateral traits. Thorax length and wing length did not exhibit consistent seasonal trends whereas wing loading significantly decreased. Significant seasonal changes in FA were not detected in any trait examined. Two traits showed reduced variation in autumn. The use of FA as an indicator of ecological stresses in insects is discussed on the basis of these results and the literature evidence.     

Does fluctuating asymmetry constitute a sensitive biomarker of nutritional stress in house sparrows (Passer domesticus)?

Small random deviations from left–right symmetry in bilateral traits, termed fluctuating asymmetry (FA), are theoretically predicted to increase with environmental stress and believed to constitute a potential biomarker in conservation. However, reported relationships between FA and stress are generally weak and variable among organisms, traits and stresses. Here we test if, and to what extent, FA increases with nutritional stress, estimated from independent feather growth measurements, in free-ranging house sparrows (Passer domesticus). Ptilochronological feather marks showed significant heterogeneity among study plots, indicating that house sparrow populations were exposed to variable levels of nutritional stress during development. However, individuals from more stressed populations did not show increased levels of fluctuating asymmetry in tarsus or rectrix length, nor was there evidence for significant between-trait concordance in FA at the individual or the population level. Lack of support for FA in tarsus and rectrix length as estimator of nutritional stress in house sparrows may indicate that developmental instability is insensitive to nutritional stress in this species, poorly reflected in patterns of fluctuating asymmetry due to ecological or statistical reasons, or highly context-specific. Such uncertainty continues to hamper the use of FA as a biomarker tool in conservation planning.     

Dental arch asymmetry in an isolated Adriatic community

Developmental stability reflects the ability of a genotype to develop in the same way under varying environmental conditions. Deviations from developmental stability, arising from disruptive effects of environmental and genetic stresses, can be measured in terms of fluctuating asymmetry, a particularly sensitive indicator of the ability to cope with these stresses during ontogeny. In an inbred Adriatic island population, we expected dental arch fluctuating asymmetry 1) to be higher than in an outbred sample from the same island, and 2) within this population, to increase with the level of inbreeding. Due to environmental stress, we also expected to find higher fluctuating asymmetry in the outbred island population than in an urban reference group from the same country. The material consisted of 506 dental casts of 253 children from 1) the island of Hvar, and 2) Zagreb, Croatia. Three-dimensional coordinates of 26 landmarks spanning the arches were digitized. The analysis partitioned the asymmetry of arch forms into components for directional and fluctuating bilateral asymmetry, using the appropriate Procrustes method (geometric morphometrics). The results corroborated the hypotheses. Fluctuating asymmetry was found to be higher on the island than in Zagreb in all groups and in both jaws, and increased significantly with endogamy level in the lower jaw. There was no significant directional asymmetry in the Zagreb sample and likewise none in the upper jaws of the outbred island group, but significant directional asymmetry in both jaws of the inbred population and also in the lower jaws of the outbred island group. These results suggest an environmental as well as a genetic influence on dental arch asymmetry. Although the lower jaws expressed these two stresses almost additively, the upper jaws appeared to be better buffered. The role of directional asymmetry as a potential indicator of craniofacial developmental instability clearly merits further attention.