精确重建古大气CO2浓度的气孔方法

为了更好地理解大气CO2浓度与气候之间的耦合关系,维护自然与人类社会的可持续发展,精确重建古大气CO2浓度的变化尤为重要。在众多研究古大气CO2浓度的方法中,化石植物叶片气孔方法已被证明是一种比较精确和理想的方法。文章介绍了气孔方法恢复古大气CO2浓度的概念、方法以及最新应用,用气孔比率法恢复了中国西北地区早、中侏罗世的大气CO2浓度变化。其趋势为：从早侏罗世普林斯巴赫期一中侏罗世阿林期一巴柔期,大气CO2浓度先是下降,而后持续上升,整个时期的大气CO2浓度在1000—1500ppmv之间,这种趋势与Bernet的碳平衡模型基本一致,并指出恢复的古大气CO2浓度低于碳平衡模型值可能是由于海拔引起的。最后提出了几个方面的研究前景。     

华北低丘山区栓皮栎人工林冠层CO2浓度和δ13C变化及其影响因素

采用离轴积分腔输出光谱技术测定华北低丘山区栓皮栎人工林冠层上缘(11 m)和下部(6 m)大气CO2浓度和δ13C值,在小时尺度上分析冠层CO2浓度和δ13C变化及其影响因素.结果表明:冠层CO2浓度呈先高后低再升高的日变化趋势,而δ13C值没有明显一致的日变化规律.白天大气不稳定状态出现的频率为70.2%,在光合作用和林内湍流的共同作用下,栓皮栎冠层下部CO2浓度高于冠层上缘约1.70μmol·mol-1,而δ13C值低于冠层上缘约0.81‰.晚上大气稳定状态出现的频率为76.2%,湍流弱,冠层叶片呼出的CO2不易流动,导致冠层下部CO2浓度高于上缘约1.24μmol·mol-1,δ13C值低于冠层上缘约0.58‰.白天和晚上冠层上下缘的CO2浓度差值与δ13C差值均呈显著的相关关系.逐步回归分析表明,白天太阳辐射和相对湿度是影响冠层CO2浓度和δ13C值差异的主要环境因子,晚上温度显著影响冠层下部与上缘δ13C值的变化,这些环境因子通过增强或减弱光合和呼吸作用来影响冠层大气中CO2浓度和δ13C值的变化.     

施氮和大气CO2浓度升高对小麦旗叶光合电子传递和分配的影响

采用开顶式气室盆栽培养小麦,设计2个大气CO2浓度(正常:400 μmol.mol-1;高:760 μmol·mol-1)、2个氮素水平(0和200 mg·kg-1土)的组合处理,通过测定小麦抽穗期旗叶氮素和叶绿素浓度、光合速率(Pn)-胞间CO2浓度(C1)响应曲线及荧光动力学参数,来测算小麦叶片光合电子传递速率等,研究了高大气CO2浓度下施氮对小麦旗叶光合能量分配的影响.结果表明:与正常大气CO2浓度相比,高大气CO2浓度下小麦叶片氮浓度和叶绿素浓度降低,高氮处理的小麦叶片叶绿素a/b升高.施氮后小麦叶片PSⅡ最大光化学效率(Fv/Fm)、PSⅡ反应中心最大量子产额(Fv'/Fm')、PSⅡ反应中心的开放比例(qr)和PSⅡ反应中心实际光化学效率(φPSⅡ)在大气CO2浓度升高后无明显变化,虽然叶片非光化学猝灭系数(NPQ)显著降低,但PSⅡ总电子传递速率(JF)无明显增加;不施氮处理的Fv'/Fm'、φPSⅡ和NPQ在高大气CO2浓度下显著降低,尽管Fv/Fm和qp无明显变化,JF仍显著下降.施氮后小麦叶片JF增加,参与光化学反应的非环式电子流传递速率(Jc)明显升高.大气CO2浓度升高使参与光呼吸的非环式电子流传递速率(J0)、Rubisco氧化速率(V0)、光合电子的光呼吸/光化学传递速率比(J0/Jc)和Rubisco氧化/羧化比(V0/Vc)降低,但使Jc和Rubisco羧化速率(Vc)增加.因此,高大气CO2浓度下小麦叶片氮浓度和叶绿素浓度降低,而增施氮素使通过PSⅡ反应中心的电子流速率显著增加,促进了光合电子流向光化学方向的传递,使更多的电子进入Rubisco羧化过程,Pn显著升高.     

大气CO_2浓度升高对香蕉光合作用及光合碳循环过程中叶氮分配的影响

生长在高CO2 浓度 (70 0± 5 6 μl·L-1) 1周的香蕉叶片 ,其光合速率 (Pn ,μmol·m-2 ·s-1)为 5 .14± 0 .32 ,较生长在大气CO2 浓度 (35 6± 30lμl·L-1)的高 2 2 .1% ,而生长在较高CO2 浓度下 8周 ,叶片Pn较生长在大气CO2 浓度的低 18.1% ,表现香蕉叶片对较长期高CO2 浓度的驯化和光合作用抑制 .生长在高CO2 浓度的香蕉叶片有较低光下呼吸速率 (Rd) ,而不包括光下呼吸的CO2 补偿点则变幅较小 .最大羧化速率 (Vcmax)和电子传递速率 (J)分别较生长在大气CO2 浓度的低 30 .5 %和 14 .8% ,根据气体交换速率计算的表观量子产率 (α ,molCO2·mol-1光量子 ) ,生长在较高CO2 浓度下 8周的叶片为 0 .0 14± 0 .0 1,而生长在大气CO2 浓度下的为 0 .0 2 5±0 .0 0 5 .较高CO2 浓度下叶片的表观量子产率降低 44% .光能转换效率 (electrons·quanta-1)亦从 0 .2 0 3降低至0 .136 .生长在较高CO2 浓度下香蕉叶片的叶氮在Rubicos分配系数 (PR)、叶氮在生物力能学组分分配系数(PB)和叶氮在光捕组分的分配系数 (PL)均较生长在大气CO2 浓度低 ,表明在高CO2 浓度下较长期生长 (8周 )的香蕉叶片多个光合过程受抑制 ,光合活性明显降低 .     

大气CO2升高和蚯蚓活动对土壤C、N的影响

以加倍CO2浓度（750μmol/mol）处理和正常CO2浓度（370μmol/mol）生长下的棉花凋落叶为试验材料,以威廉腔蚓Metaphire guillemi（Michaelsen,1895）和不同的CO2浓度（750μmol/mol和370μmol/mol）为作用因子,分析了蚯蚓、CO2浓度通过叶片分解对土壤C、N含量的影响。结果表明：接种蚯蚓和加入凋落叶的联合作用对有机C有显著提高作用。接种蚯蚓对土壤全N含量影响不显著,但CO2浓度升高和蚯蚓联合作用对土壤全N含量有显著影响。CO2、叶片、蚯蚓3因子联合作用对土壤C、N含量有显著提高作用,且与蚯蚓和叶片联合作用对土壤C、N含量的影响相比,其效果更显著。结果显示,CO2浓度的升高通过改变植物凋落物C含量及其营养成分,影响了其潜在的降解有效性,同时大气CO2浓度的升高影响凋落物在蚯蚓体内降解过程,从而对凋落物的有效降解产生显著影响,最终改变土壤C、N含量。     

科尔沁沙地植物成熟叶片性状与叶凋落物分解的关系

采用室内培养的方法,对科尔沁沙质草地20个主要植物种(10个单子叶植物种和10个双子叶植物种)叶凋落物的CO2释放量和释放速率进行比较,同时测定了20种植物成熟叶片的热值、镁浓度、磷浓度、氮浓度、钾浓度、碳浓度、碳氮比、氮磷比、比叶面积、干物质含量以及叶面积等11项叶片性状,分析20种植物叶凋落物的CO2释放量和释放速率与11项叶片性状的相关关系.结果表明, 20种植物的叶片性状在物种间变异范围很大,大多数指标的最大值和最小值的差异在3倍以上,而个别指标如叶镁浓度差异更大,接近9倍之多.由于本项研究中的20种植物均来自于同一样地,因此认为20种植物的自身性状和遗传特性决定了其叶片性状的变异.20种植物叶凋落物培养28d的CO2释放量平均值为(4121±1713)μg kg-1,释放量最大的是尖头叶藜(8767±177)μg kg-1干土,释放量最小的是马唐(1669±47)μg kg-1,二者相差5倍以上.但20种植物叶凋落物CO2释放速率表现相同的变化趋势,培养初期凋落物分解迅速,后期分解相对缓慢.比较分析发现,双子叶植物和单子叶植物叶凋落物CO2释放量、CO2释放速率,以及成熟叶片的叶氮浓度、叶碳浓度、叶C/N和叶干物质含量之间差异显著.相关分析揭示,20种植物成熟叶片叶氮浓度、叶碳浓度、叶C/N和叶干物质含量与叶凋落物分解过程中的CO2释放量和释放速率之间呈显著的相关关系,说明可以利用成熟叶片的N浓度、C浓度、C/N以及干物质含量间接的预测叶凋落物的分解速率.     

树木年轮 δ13C 值及其对我国北方大气CO2浓度变化的指示意义

 本文采样分析了承德市油松年轮中δ13C值自工业革命以来的变化,用以揭示我国北方大气CO2浓度的变化规律。结果表明：承德市油松年轮中的δ13C值自1810年以来平均下降了0.839‰,下降范围0.682‰～1.120‰,指示了大气CO2浓度逐渐升高的特点。δ13C值与历史时期全球大气CO2浓度之间存在显著相关关系(r＝ –0.5609,P<0.01)。应用树木年轮δ13C值与大气CO2浓度之间的关系式,推测出我国北方大气CO2浓度从工业革命以前的约278.4μmol·mol-1上升到目前的340μmol·mol-1。从而为我国的全球变化研究提供了CO2浓度历史变迁方面的证据。     

CO2浓度升高和模拟氮沉降对青川箭竹叶营养质量的影响

通过研究大熊猫主食竹之一的青川箭竹(Fargesia rufa Yi)叶营养质量对CO2浓度升高和模拟氮沉降的响应,预测在气候变化下箭竹和大熊猫之间的取食关系,以期为大熊猫的长久保护提供科学参考.利用人工环境控制生长室对青川箭竹幼苗进行了1个生长季节的大气CO2浓度和施氮处理:(1)CON(对照,不添加N和环境CO2浓度),(2)EC(环境CO2浓度+350μmol/mol、不添加N),(3)EN(添加5gNm-2a-1、环境CO2浓度),(4)ECN(环境CO2浓度+350 μmol/mol、添加5 g N m-2 a-1).结果表明:EC处理对青川箭竹叶片的C含量无显著影响,但降低了叶片中N和P含量,从而导致C:N增加,而N:P没显著变化.另外,EC处理增加了叶片中可溶性蛋白、可溶性糖、淀粉、蔗糖和果糖的含量,但降低了木质素和纤维素含量.同时,EC也明显增加了叶片中防御物质单宁的含量.另一方面,EN处理显著降低了叶片中C的含量,并增加了N的含量,但没有改变P的含量,从而C:N减小,而N:P增加.EN显著提高了箭竹叶片可溶性蛋白、可溶性糖和木质素含量,减少了淀粉和纤维素,但对单宁无明显影响.ECN减少了箭竹叶的单宁和N、P的含量,但显著增加了叶的可溶性蛋白、可溶性糖、果糖、蔗糖和淀粉含量.大气CO2浓度升高和氮沉降对叶的N、单宁、可溶性糖和淀粉含量具有显著的交互作用.在未来气候变化情景下,箭竹叶营养质量提高将可能影响叶的生物化学过程以及箭竹-大熊猫之间的取食关系.     

大气CO2浓度升高对绿豆叶片光合作用及叶绿素荧光参数的影响

利用FACE系统在大田条件下通过盆栽试验研究了大气CO2浓度升高[CO2浓度平均为(550+60) μmol·mo1-1]对绿豆叶片光合生理和叶绿素荧光参数的影响.结果表明:与对照[ CO2浓度平均为(389+40) μmol·mol-1左右]相比,大气CO2浓度升高使花荚期绿豆叶片净光合速率(Pn)和胞间CO2浓度(Ci)分别升高11.7％和9.8％,气孔导度(Gs)和蒸腾速率(Tr)分别下降32.0％和24.6％,水分利用效率(WUE)提高83.5％;在蕾期,CO2浓度升高对绿豆叶片叶绿素初始荧光(Fo)、最大荧光(Fm)、可变荧光(Fv)、Fv/Fm和Fv/Fo没有显著影响;在鼓粒期,CO2浓度升高使绿豆叶片Fo增加19.1％,Fm和Fv分别下降9.0％和14.3％,Fv/Fo和Fv/Fm分别下降25.8％和6.2％.表明大气CO2浓度升高可能使绿豆生长后期光系统Ⅱ反应中心结构受到破坏,叶片的光合能力下降.     

叶片碳同位素对城市大气污染的指示作用

沿南京市市中心-郊区-城市森林的梯度,观测了近地层大气CO2(pN-S(CO2))、SO2(pN-S(SO2))、NO2(pN-S(NO2))浓度,以及公路两侧悬铃木(Platanus acerifolia)叶片重金属Pb、Cr含量、稳定碳同位素13C和放射性碳同位素14C的变化规律,探讨叶片13C和14C对城市大气污染的指示作用。结果表明,沿市中心-郊区-城市森林梯度,pN-S(CO2)、pN-S(SO2)和pN-S(NO2)均呈现下降的趋势,与观测点距市中心距离间均存在极显著的负相关关系;与城市森林相比,市中心pN-S(CO2)、pN-S(SO2)和pN-S(NO2)分别高7.3%、88%和210%。沿市中心-郊区-城市森林梯度,叶片δ13C和14C含量有升高的趋势,其中,δ13范围为-27.9‰—-31.4‰,14C pMC范围为95.8%—101.2%,即叶片C有9.3%—14.4%来源于化石燃料;叶片14C含量与距市中心距离间存在极显著的线性相关。叶片1δ3C与污染参数pN-S(CO2)、pN-S(SO2)、pN-S(NO2)、Pb、Cr间均表现为负相关,其中与pN-S(CO2)、Pb间的相关性达到显著水平;而叶片14C含量与pN-S(CO2)、pN-S(SO2)、pN-S(NO2)、Pb和Cr间均表现出显著的负相关。表明,叶片14C含量可用于指示城市大气污染状况,且与叶片δ13C相比更为稳定。     

ACTION OF FUSICOCCIN ON THE POTASSIUM BALANCE OF GUARD CELLS OF PHASEOLUS VULGARIS

Fusicoccin induces stomatal opening in both the light and dark. The stomatal aperture and K content of guard cells was measured to determine whether the action of fusicoccin in inducing stomatal opening is directly related to the uptake of K by the guard cells. Both detached and attached epidermis was treated with fusicoccin and the K content was determined by staining with cobalt sodium nitrite or by electron probe microanalysis. The K content of guard cells in detached epidermal strips floated on 10 μm fusicoccin in 10 mm KCl and aqueous CH₃OH (0.02%, v/v) increased in the light and dark as the stomata opened. After exposure to fusicoccin for 6 hr in the light, however, the stomata were closed and no K could be detected in the guard cells. The K content of guard cells of attached epidermis painted with fusicoccin also increased as the stomata opened, but the concentration of K in the subsidiary cells was not significantly altered by fusicoccin-stimulated opening. Moreover, painting with fusicoccin did not significantly change the Ca and P content of the guard or subsidiary cells. Stomata of epidermal strips, opened to their maximum width by fusicoccin, showed only a small and temporary closure when transferred to a solution of 10 μm abscisic acid. The use of metabolic inhibitors suggested that energy for the uptake of the K may be provided by both photophosphorylation and oxidative phosphorylation.     

中国桑寄生科植物叶表皮微形态

通过扫描电镜对中国桑寄生科桑寄生亚科8属18种和槲寄生亚科1属2种植物成熟叶的上、下叶表皮内表面和下表皮外表面进行了研究。内面观发现桑寄生科植物叶上、下表皮形状为多边形,垂周壁式样平直或稍弓形,常具有角质增厚,平周壁常覆盖厚角质或颗粒状、丝状角质增厚;气孔存在于上下表皮,通常下表皮较多,气孔的形状,特别是保卫细胞的形态在亚科间、属间或种间都具有一定的差异,气孔器类型为平列型或单圈型。下表皮表面观察了的角质膜和蜡质纹饰、气孔的形状,外部气孔缘及外部气孔缘内缘的特征。这些特征在亚科或属级水平上较为稳定,有的也表现出种间差异,有一定的分类价值。从气孔形态和外部气孔周围角质膜来看,两亚科显示出明显的不同:桑寄生亚科上、下表皮均具有内部气孔缘,而槲寄生亚科没有此结构;桑寄生亚科外部气孔周围角质膜增厚成环状,其上具增厚的条纹,而槲寄生亚科外部气孔周围角质膜增厚成脊状,不具条纹。这些特征支持槲寄生亚科作为独立1个科来处理。     

盾叶薯蓣四倍体与二倍体叶表皮及气孔器显微结构的比较

对盾叶薯蓣四倍体和二倍体阴叶和阳叶表皮及气孔器的显微形态结构进行了比较。结果显示:叶片表皮细胞及气孔器的形态结构因染色体倍性及光照的不同而存在明显差异。表皮细胞有规则多边形与不规则细胞壁内褶2种类型;气孔器为典型的双子叶植物无规则型气孔器,仅分布在下表皮。四倍体表皮细胞密度、气孔器密度及气孔指数平均值分别为476.28±6.87个.mm-2、78.22±3.1个.mm-2、14.11,较二倍体的分别小4.60%、17.95%和11.98%。前者气孔器长、宽及保卫细胞宽度平均值分别为32.78±2.09μm、26.07±1.55μm、9.63±1.14μm,较后者的分别大10.73%、3.90%和18.01%,差异极显著。前者阴叶的较其阳叶的分别大5.77%、6.00%、8.72%;后者的则分别大1.72%、1.74%、2.41%。前者叶片保卫细胞中的叶绿体数目为23.93±3.19个,较后者的多74.00%。表皮细胞及气孔器密度、气孔指数,气孔器大小,保卫细胞叶绿体数目、尤其是保卫细胞宽度,可作为倍性鉴定的参考依据。四倍体高产有其良好的叶片结构基础。     

The CO(2) response of Vicia guard cells acclimates to growth environment

Stomata of growth chamber-grown Vicia faba leaves have an enhanced CO(2) response, measured as change in stomatal aperture, compared to stomata of greenhouse-grown leaves. Reciprocal transfer experiments showed that the stomatal response to CO(2) acclimated to the growing environment. Stomata of growth chamber-grown leaves transferred to a greenhouse lost their high CO(2) sensitivity within 2-3 d while stomata of greenhouse-grown leaves transferred to a growth chamber acquired a high CO(2) sensitivity within 5-7 d. Experiments measuring the CO(2) responses of stomata in detached epidermis showed that growth chamber and greenhouse-grown stomata have the same contrasting CO(2) sensitivity observed in the intact leaf, indicating that the responses reflect intrinsic guard cell properties. The acclimation properties of the CO(2) response of guard cells have implications for the understanding of stomatal function under the predicted increases in atmospheric CO(2).     

The too many mouths and four lips mutations affect stomatal production in Arabidopsis

Stomata regulate gas exchange through the aerial plant epidermis by controlling the width of a pore bordered by two guard cells. Little is known about the genes that regulate stomatal development. We screened cotyledons from ethyl methanesulfonate-mutagenized seeds of Arabidopsis by light microscopy to identify mutants with altered stomatal morphology. Two mutants, designated too many mouths (tmm) and four lips (flp), were isolated with extra adjacent stomata. The tmm mutation results in stomatal clustering and increased precursor cell formation in cotyledons and a virtual absence of stomata in the inflorescence stem. The flp mutation results in many paired stomata and a small percentage of unpaired guard cells in cotyledons. The double mutant (tmm flp) exhibits aspects of both parental phenotypes. Both mutations appear to affect stomatal production more than patterning or differentiation. tmm regulates stomatal production by controlling the formation, and probably the activity, of the stomatal precursor cell.     

Stomatal movement and potassium transport in epidermal strips of Zea mays: The effect of CO2

Summary The correlation between stomatal action and potassium movement in the epidermis of Zea mays was examined in isolated epidermal strips floated on distilled water. Stomatal opening in the isolated epidermis is reversible in response to alternate periods of light or darkness, and is always correlated with a shift in the potassium content of the guard cells. K accumulates in guard cells during stomatal opening, and moves from the guard cells into the subsidiary cells during rapid stomatal closure. When epidermal strips are illuminated in normal air, as against CO₂-free air, the stomata do not open and there is a virtually complete depletion of K from the stomatal apparatus. In darkness CO₂-containing air inhibits stomatal opening and K accumulation in guard cells, but does not lead to a depletion of K from the stomata as observed in the light.     

The pattern of cuticular blue-fluorescing phenolics deposition during the development of Prunus persica leaves

Although stomatal ontogeny is closely related to the development and maturation of the epidermal tissue, stomatal patterns in relation to cuticle construction and cuticular material deposition during leaf development have not received adequate attention. We observed the deposition of blue-fluorescing cuticular phenolics over guard and epidermal cells, as well as stomatal formation and patterning using the alkali-induced blue fluorescence of the cuticle of Prunus persica leaves. Stomata of different stages of maturity occurred together during leaf development, mainly at the tip of the lamina. The deposition of fluorescing compounds initially appeared over the guard cells of the developing stomata complexes and gradually extended to the neighbouring epidermal cells. Based on the blue fluorescence emitted by the cuticular layers, we constructed digital maps of leaves of different developmental stages, showing the pattern of stomatal formation and deposition of fluorescing compounds. A longitudinal tip-to-base gradient in the formation of stomata, as well as in the deposition of fluorescing compounds was observed in young developing leaves. The deposition of blue-fluorescing phenolic compounds seems to be coordinated with stomatal development.     

三种国产阴地蕨科植物叶的表皮构造

运用光学显微镜和扫描电镜对三种国产阴地蕨科植物：劲直蕨萁(Botrypus strictus)、小阴地蕨(Botrychium lunaria)和薄叶阴地蕨(Sceptridium daucifolium)〖WTBZ〗叶的成熟表皮构造进行了详细观察和研究。它们具有共同的特征：气孔散生,气孔类型无规则型,气孔长轴方向多与叶脉延伸方向一致。但也存在明显区别,特别是劲直蕨萁与另外二个种的区别更为明显：前者表皮细胞垂周壁直,相邻气孔不接触,保卫细胞平周壁具细条纹。但后二者之间亦有一定区别：薄叶阴地蕨的气孔为下生式,不下陷,而小阴地蕨的气孔为两面气孔型,气孔下陷。本文是国内首次对阴地蕨科叶的表皮构造进行研究,其研究结果表明,表皮构造在阴地蕨科植物的鉴定上具有重要意义,并且在研究阴地蕨科的分类以及起源和演化上也有一定参考价值。     

Rapid low temperature-induced stomatal closure occurs in cold-tolerant Commelina communis leaves but not in cold-sensitive tobacco leaves, via a mechanism that involves apoplastic calcium but not abscisic acid

Commelina communis stomata closed within 1 h of transferring intact plants from 27 degrees C to 7 degrees C, whereas tobacco (Nicotiana rustica) stomata did not until the leaves wilted. Abscisic acid (ABA) did not mediate cold-induced C. communis stomatal closure: At low temperatures, bulk leaf ABA did not increase; ABA did not preferentially accumulate in the epidermis; its flux into detached leaves was lower; its release from isolated epidermis was not greater; and stomata in epidermal strips were less sensitive to exogenous ABA. Stomata of both species in epidermal strips on large volumes of cold KCl failed to close unless calcium was supplied. Therefore, the following cannot be triggers for cold-induced stomatal closure in C. communis: direct effects of temperature on guard or epidermal cells, long-distance signals, and effects of temperature on photosynthesis. Low temperature increased stomatal sensitivity to external CaCl(2) by 50% in C. communis but only by 20% in tobacco. C. communis stomata were 300- to 1,000-fold more sensitive to calcium at low temperature than tobacco stomata, but tobacco epidermis only released 13.6-fold more calcium into bathing solutions than C. communis. Stomata in C. communis epidermis incubated on ever-decreasing volumes of cold calcium-free KCl closed on the lowest volume (0.2 cm(3)) because the epidermal apoplast contained enough calcium to mediate closure if this was not over diluted. We propose that the basis of cold-induced stomatal closure exhibited by intact C. communis leaves is increased apoplastic calcium uptake by guard cells. Such responses do not occur in chill-sensitive tobacco leaves.     

不同产地何首乌叶表皮结构的解剖学特征与气候因子的关系

在光学显微镜下观察了不同产地何首乌叶表皮结构特征,应用多元回归方法对不同产地何首乌的叶表皮特征与气候因子的关系进行了分析。观察的叶表皮特征指标有气孔密度、气孔指数、气孔器长、气孔器宽、气孔极区角质加厚、气孔器类型、表皮细胞垂周壁性状及叶表面角质条纹。观察结果:上表皮有少量气孔器分布;在下表皮,气孔器类型为非典型不等细胞型和不规则型,有少量腺鳞分布,气孔器密度每1mm2为241.7(64～573)个,气孔指数为17.1(7.5～26.5)%,气孔器长31.1(20～44)μm,气孔器宽23.1(16～38)μm。随着产地的不同,何首乌叶下表皮结构有明显差异。分析结果显示气孔器、气孔器宽度以及气孔密度均与纬度关系密切,随纬度的升高,气孔器长、气孔器宽呈减小的趋势,气孔密度呈增加的趋势,R2值分别为0.619、0.729、0.772。