蚜虫取食中的细胞壁修饰与免疫功能

蚜虫是世界性害虫,它通过独特的口针结构和丰富的唾液组分破坏植物细胞壁,穿过表皮细胞和叶肉细胞间隙,克服多重植物抗性,到达韧皮部取食为害。已有报道蚜虫唾液中含有多种细胞壁修饰酶能够降解修饰细胞壁,帮助蚜虫在细胞间刺探,更为有效的定位韧皮部。而细胞壁作为保护植物细胞的重要屏障,能感知和传递细胞壁损伤信号,通过调控细胞壁修饰酶的表达水平启动胞内诱导抗性,从而影响蚜虫的刺探、取食和定殖。此外,蚜虫唾液中的一些效应因子还能抑制细胞壁免疫和胞内抗性。可见,细胞壁免疫在蚜虫持续取食和成功定殖中发挥重要功能。为深入理解细胞壁免疫在蚜虫刺探与取食过程中的作用机制,本文概述了蚜虫唾液关键组分对细胞壁修饰与免疫的调控作用,从植物细胞壁多糖结构修饰、损伤信号传导和胞内抗性等方面重点论述对蚜虫取食行为的影响,结合病原菌与细胞壁免疫互作机制,进一步揭示蚜虫与细胞壁免疫互作新机制,为基于阻断蚜虫韧皮部取食的分子抗虫育种提供新思路。     

植物和刺吸式口器昆虫的诱导防御与反防御研究进展

刺吸式口器昆虫在长期的进化过程中形成特殊的口针结构,用于专门吸食植物韧皮部筛管细胞的汁液成分.以蚜虫为例,它们在取食过程中分泌的胶状唾液和水状唾液将有效的降低植物防御反应,其中水状唾液包含的大量酶类不仅可以帮助蚜虫穿刺植物韧皮部,刺探到筛管细胞,同时也是植物感受蚜虫为害的激发因子,诱导出植物防御反应和相关抗性基因的表达...     

电子取食监测仪在植物抗虫性研究中的应用

主要综述了电子取食监测仪在植物对同翅目昆虫抗虫性研究中的应用情况.利用电子取食监测仪可监测同翅目昆虫（蚜虫、飞虱、叶蝉、蓟马、粉虱等）在不同抗、感虫品种上的取食行为,并根据取食行为参数如刺探频率、唾液分泌时间、韧皮部取食时间等在不同抗、感品种上的差异来区别品种的抗、感虫特性和确定抗虫因子存在的部位.     

昆虫对植物次生物质的代谢适应机制及其对昆虫抗药性的意义

植物次生物质(plant secondary metabolites)对昆虫的取食行为、生长发育及繁殖可以产生不利影响,甚至对昆虫可以产生毒杀作用。为了应对植物次生物质的不利影响,昆虫通过对植物次生物质忌避取食、解毒代谢等多种机制,而对寄主植物产生适应性。其中,昆虫的解毒代谢酶包括昆虫细胞色素P450酶系（P450s）及谷胱甘肽硫转移酶（GSTs）等,在昆虫对植物次生物质的解毒代谢及对寄主植物的适应性中发挥了重要作用。昆虫的解毒酶系统不仅可以代谢植物次生物质,还可能代谢化学杀虫剂,因而昆虫对寄主植物的适应性与其对杀虫剂的耐药性甚至抗药性密切相关。昆虫细胞色素P450s和GSTs等代谢解毒酶活性及相关基因的表达可以被植物次生物质影响,这不仅使昆虫对寄主植物的防御产生了适应性,还影响了昆虫对杀虫剂的解毒代谢,因而改变昆虫的耐药性或抗药性。掌握昆虫对植物次生物质的代谢适应机制及其在昆虫抗药性中的作用,对于明确昆虫的抗药性机制具有重要的参考意义。本文综述了植物次生物质对昆虫的影响、昆虫对寄主植物次生物质的代谢机制、昆虫对植物次生物质的代谢适应性对昆虫耐药性及抗药性的影响等方面的研究进展。     

E型麦二叉蚜在不同抗性表型燕麦上的取食行为

【目的】为了明确E型麦二叉蚜Schizaphis graminum在不同抗性表型燕麦品系上刺探取食行为的差异性,查明燕麦抗蚜因子。【方法】采用直流型刺吸电位仪(DC-EPG Giga-4)记录了E型麦二叉蚜无翅成蚜在10个抗性表型差异MN品系皮燕麦上A,B,C,Pd,E,F,G和np 8种基本波形的发生次数和持续时间。【结果】刺探电位图谱显示,E型麦二叉蚜在10个抗性表型差异MN品系皮燕麦上的识别时间在1.03~2.62 min之间。表型中抗的MN10253品系上非刺探时间相对较长,开始至首次刺探时间最长,分别为117.77和13.80 min,说明叶片表面可能存在不利于E型麦二叉蚜定位和识别寄主的干扰因子;表型中抗的MN08252品系上口针在叶肉组织和木质部刺探取食持续总时间以及开始至首次刺探木质部时间均为最短,分别为78.01,62.73和32.29 min,且np波发生次数多达28.57次,说明叶肉组织和木质部中可能存在抗蚜因子;表型低抗的MN06203品系上F波持续总时间和平均持续时间均最长,分别为4.96 min和5.81 s,说明该品系具有较强的机械抗性;表型低感的MN11213品系上E1时间最短,为1.94 min,开始至首次刺探韧皮部的时间最长,为246.05 min,说明抗蚜因子可能存在于韧皮部。【结论】皮燕麦表型中抗的MN10253品系叶片表面、MN08252品系的叶肉组织和木质部以及表型低感的MN11213品系韧皮部组织内可能存在抗蚜性因子,表型低抗的MN06203品系机械抗性较强。     

转换寄主前后扶桑绵粉蚧取食行为的EPG分析

【目的】分析扶桑绵粉蚧Phenacoccus solenopsis在发生寄主转移前后的取食行为,以据此评价其寄主适应性。【方法】利用刺吸电位（EPG）技术,量化扶桑绵粉蚧转换寄主植物前后的取食行为并进行对比分析。【结果】寄主转换后,扶桑绵粉蚧的取食行为受到显著的影响,需要耗费更多的时间到达韧皮部,总取食时间显著变短,取食效率降低。但寄主转换后,下一代成虫的取食效率提高,总刺探数、取食刺探数、总刺探时间以及到达韧皮部的效率与时间均与寄主转换前的扶桑绵粉蚧个体不存在显著差异。【结论】扶桑绵粉蚧的取食行为存在高度可塑性,这种特性利于扩大其寄主植物范围,有可能是促其成为重要农业入侵生物的原因之一。     

温室白粉虱取食行为的刺探电位(EPG)研究

该文揭示了温室白粉虱Trialeurodes vaporariorum Westwood成虫及幼虫的刺探电位波形与其刺探、取食、产卵行为之间的对应关系,并讨论了这项技术在研究植物抗虫机理方面的应用价值。在成虫,A波、C波分别代表刺探的开始和进行过程;F波代表刺探过程中遇到机械障碍;G波表示吸食木质部导管汁液;E(pd)的(1)和(2)分别表示与取食韧皮部筛管汁液有关的两种行为;粉虱的产卵波形分为两种亚波,分别由Ovi-I和Ovi-II表示,各自代表产卵时的两种行为：产卵器接触并划破叶表皮及卵柄插入叶组织。在幼虫,H波代表吸食筛管液,而L波则表示在筛管细胞内的一种非吸食行为。幼虫蜕皮时先拔出口针,新龄期的幼虫将其口针重新刺入叶组织。     

番茄黄化曲叶病毒对烟粉虱MED隐种取食行为的影响

为探究番茄黄化曲叶病毒（TYLCV）对烟粉虱刺探取食行为的影响并研究其与烟粉虱之间的互作关系。利用EPG记录了感染TYLCV的MED隐种在番茄寄主上的取食波形,并与未感染TYLCV的MED隐种的取食波形进行了比较研究。结果表明,烟粉虱开始取食行为后,带毒MED和未带毒MED平均刺探次数和非刺吸时间占总取食时间的比例方面没有显著差异。带毒MED从口针刺探开始至到达韧皮部的平均时间为46 min,从记录开始至口针第一次到达韧皮部的平均时间为135 min,未带毒MED相对应的平均时间分别为19 min和25 min,两者结果差异显著;在烟粉虱取食番茄韧皮部阶段,带毒和未带毒MED的唾液分泌总时间分别为9 min、3 min,第一次韧皮部持续吸食平均时间分别为143 min、26 min,带毒MED显著高于未带毒MED。带毒MED在第一次持续吸食至第一次取食结束的平均时间为15 min,显著高于未带毒MED的4 min。综上所述,带毒MED完成整个刺探取食行为的平均时间较未带毒MED显著增加,特别是唾液的分泌时间以及第一次在韧皮部的持续吸食时间。表明,TYLCV能促进MED加强唾液分泌和韧皮部取食,从而促进病毒的传播。     

亚致死剂量溴氰虫酰胺对MED烟粉虱取食行为及传播TYLCV的影响

取食是烟粉虱Bemisia tabaci(Gennadius)传播植物病毒最重要的途径之一.本研究利用昆虫刺探电位(EPG)技术研究了杀虫剂溴氰虫酰胺亚致死浓度(LC15)对携带番茄黄化曲叶病毒(TYLCV)MED烟粉虱取食行为和传毒效率的影响.结果表明,溴氰虫酰胺对未携毒和携毒MED烟粉虱的亚致死浓度(LC15)分别为1.52 mg/kg和1.75 mg/kg,差异不显著.携毒烟粉虱在溴氰虫酰胺亚致死浓度处理的番茄苗上取食,第一次刺探起始时间显著推迟,是对照番茄苗上的3.7倍.在韧皮部取食阶段,处理组烟粉虱在韧皮部分泌唾液平均时间、吸食汁液总时间和平均时间均显著缩短,分别是对照组的0.10倍、0.14倍和0.10倍,韧皮部取食能力显著降低.qPCR试验研究表明,溴氰虫酰胺亚致死浓度处理极显著的降低了烟粉虱的传毒效率,携毒烟粉虱取食48 h的番茄苗培养10 d后,TYLCV相对含量仍接近于0,表明溴氰虫酰胺对烟粉虱传播TYLCV具有良好的控制潜力.研究结果为科学合理的使用化学农药防治烟粉虱及其传播的双生病毒病提供理论依据.     

园艺矿物油乳剂对亚洲柑橘木虱成虫刺吸行为的影响

目的】亚洲柑橘木虱Diaphorina citri是传播柑橘最重要病害黄龙病(huanglongbing, Candidatus Liberibacter asiaticus)的媒介昆虫,树冠喷施园艺矿物油(horticultural mineral oil, HMO)可以减少其在柑橘上的取食和产卵。本研究旨在探索植物组织内残留的园艺矿物油如何影响亚洲柑橘木虱成虫的取食行为。【方法】采用直流型刺吸电位仪(DC-EPG Giga-8)记录亚洲柑橘木虱成虫12 h内在喷施不同浓度(0.25%, 0.5%, 1%和2%, v/v)HMO(nC24)乳剂和清水(对照)时柑橘嫩叶上的刺探和取食行为并转换为波形信号,分析比较各处理的波形参数。【结果】不同浓度HMO乳剂处理显著提高了亚洲柑橘木虱成虫在柑橘叶片上的C波(路径波)持续时间百分比,显著减少了E2波(韧皮部吸食汁液)持续时间百分比,而对D(口针第一次接触韧皮部组织), E1(韧皮部出现唾液分泌) 和G(木质部吸食汁液) 波持续时间百分比没有显著影响。HMO处理组C波的每成虫波形累积时间(WDI)显著长于对照组,而各浓度处理之间没有显著差异,但D波WDI明显短于对照组。随着HMO浓度的增加, E1, E2和G波WDI显著下降,2% HMO处理组E1, E2和G波WDI明显低于其他浓度处理组。HMO乳剂处理明显减少了刺探次数;在HMO处理中,每成虫刺探次数和每成虫各波形次数均显著少于对照组,其中以1%和2% HMO乳剂处理最少。HMO乳剂处理组np波 (非刺探波)、D波、E2波的每成虫单次波形持续时间(WDE)均短于对照组,其中2% HMO乳剂处理后,D, E1和E2波WDE显著短于其他浓度处理和对照。同时,2% HMO乳剂处理推迟了从口器接触叶片到第1次开始刺探的时间和从口器接触叶片到第1次刺探到韧皮部的时间。【结论】结果表明,2% HMO乳剂显著减少亚洲柑橘木虱成虫在柑橘叶片上的取食次数和有效取食时间,同时还缩短了其在韧皮部分泌唾液和吸食汁液时间,因此可推荐矿物油用于柑橘黄龙病和亚洲柑橘木虱的综合防治体系。同时初步推断矿物油的作用机制可能是其进入植物气孔阻止植物挥发性物质的释放,抑制或掩盖柑橘叶片表面的挥发性物质从而减少了亚洲柑橘木虱在寄主上取食。     

CELLULAR AND CHEMICAL SAMPLING DURING PHLOEM FINDING AND HOST-PLANT ACCEPTANCE BY HOMOPTERAN INSECTS

Abstract The chemical compounds that exist in plant leaf tissue are of great importance for homopteran insects during the proccess of host-plant acceptance. From the leaf surface to the phloem tube, various groups of chemicals can be detected by insect's receptors which lead to different feeding responses. The effects of sugars and alkaloids in leaf surface; polysaccharides, pH gradients and phenolics in mesophyll; amino acids, proteins, carbohydrates and secondary substances in phloem tube on phloem finding and host-plant acceptance are reviewed respectively in this paper, which also includes the potential effects of mesophyll structures on insect's feeding preference. Furthermore the mechanism by which the insects perceive these chemical and mechanical cues, correlated with the EPG (electrical penetration graph) patterns (a series of visible signals showing the stylet activities in plant tissue) is also discussed.     

Analysis of resistance in tomato and sweet pepper against the greenhouse whitefly using electrically monitored and visually observed probing and feeding behaviour

Different types of plant resistance against the greenhouse whitefly, Trialeurodes vaporariorum, were distinguished according to the tissue location of the resistance factors. The effects of resistance factors were compared by measuring the electrically monitored and visually observed probing and feeding behaviour of whiteflies on two resistant tomato genotypes (82207 and 82216), a susceptible tomato cultivar, and a non-host plant, sweet pepper. On sweet pepper, whiteflies displayed very short first probes, very long pathway probing and spent much time on non-feeding activities such as walking and standing still. Also, a high percentage of whiteflies rejected sweet pepper without ingesting substances from the phloem vessel. These data suggest a strong resistance that is based on the factors present in surface/epidermis and/or mesophyll layers of this plant. The behaviour of whiteflies on tomato 82207 was very different from that on sweet pepper. On tomato 82207 whiteflies apparently did not perceive resistance factors on the leaf surface and in the mesophyll. Resistance factors appeared to be present in the phloem tissue, because a higher number of phloem phases, longer phloem salivation periods and shorter phloem ingestion periods were observed when compared with the susceptible tomato cultivar. Tomato 82216 showed no clear resistance factor in the phloem. The importance of combined EPG and behavioural research for host-plant resistance studies are discussed.     

Plant feeding site selection on soybean by the facultatively phytophagous predator Orius insidiosus

Experiments were conducted to test whether the facultatively phytophagous predator Orius insidiosus (Say) (Heteroptera: Anthocoridae) ingested phloem, xylem or mesophyll contents from soybean plants (Glycine max L.). Potential uptake of phloem sap was examined by radiolabeling photosynthate with ¹⁴CO₂ and then measuring the accumulation of radiolabeled metabolites in feeding animals. Most O. insidiosus feeding on radiolabeled plants ingested no or very low levels of label; only 3% ingested small amounts of label, indicating the experimental insects fed very little, if at all, on the phloem. In contrast, well known phloem feeding insects used as positive controls accumulated substantial levels of labeled metabolites after feeding on known host plants. O. insidiosus did feed on xylem contents, as shown by ingestion of safranin-labeled xylem fluid. A few of the insects showed signs of feeding on the mesophyll, as indicated by the presence of chloroplasts in the gut. However, the small diameter of the food canal may cause limited passage of chloroplasts, which would contribute to an underestimation of the frequency of mesophyll feeding. Some radiolabeled metabolites remain in the mesophyll so those insects that ingested low levels of radiolabel probably ingested label from the mesophyll, which supports the notion that some level of mesophyll feeding occurred. Feeding site determines the nutrients ingested during phytophagy. These insects obtain water from the xylem, and may ingest small amounts of starches, sugars, and amino acids from the mesophyll. The results suggest that facultative phytophagy by this heteropteran predator primarily provides the insect with water, but also may provide some nutrients that supplement a prey diet and help the predator survive periods when prey are scarce.     

Maternal reproductive decisions are independent of feeding in the black bean aphid,Aphis fabae

Aphids are phloem feeders and an important assumption has been that reproduction is initiated only after phloem ingestion. Here we investigate the plant tissue location of parturition cues in winged and wingless, summer virginoparae and autumn migrants (gynoparae) of the black bean aphid, Aphis fabae. These seasonal forms have different host preferences. Using electrical penetration graph (EPG: to observe activity of the mouthparts) and video-monitoring procedures we demonstrate that the time to first parturition after host-plant contact is significantly shorter than the time to first registered phloem contact in the summer winged form. In gynoparae, the time to first parturition does not significantly differ from time to first phloem contact but is shorter than time to first phloem ingestion. Times to first parturition, first registered phloem contact and first phloem ingestion do not differ significantly in the summer wingless form. Simultaneous EPG and video recording procedures show that a high proportion of individuals of all morphs (45-70%) initiate reproduction before sustained phloem activities (salivation/ingestion). The only behaviours that all individuals demonstrate before first parturition are ‘non-penetration’ (aphid on plant surface with mouthparts outside plant) and stylet ‘pathway activity’, including secretion of gelling saliva and penetration of the non-vascular (epidermis and mesophyll) cells. A short period of penetration of the peripheral plant tissues (five cell punctures per individual) by aphids tethered and monitored by EPG decreases the time to first parturition of the winged summer form when subsequently placed on a Parafilm sachet containing 15% sucrose solution. This treatment also significantly increases the incidence of reproduction and individual reproductive output of gynoparae over a 24-h period. No detectable effects of tissue penetration on subsequent reproductive output are observed in the wingless summer form. Additionally, EPGs reveal that a number of aphids of all morphs display xylem ingestion, which occurs predominantly before initiation of phloem feeding but is not necessary to initiate parturition. It is concluded that aphids are likely to detect parturition cues during stylet punctures of cells within peripheral tissue layers, before reaching the phloem vessels.     

Behavioral Sabotage of Plant Defense: Do Vein Cuts and Trenches Reduce Insect Exposure to Exudate?

Many insect folivores sever veins or cut trenches before feeding on leaves that emit fluid from latex, resin, or phloem canals. The behaviors reportedly function to reduce the insect's exposure to exudate. This hypothesis was tested with three species that sever leaf midribs and two species that trench. In each case, the insect's cuts reduced exudation at the distal feeding site by at least 94% relative to an adjacent control leaf. However, most of the insects contacted exudate during the vein-cutting/trenching operation, which could potentially negate any benefits received during feeding. To estimate how much exudate red milkweed beetles (Tetraopes tetrophthalmus) ingest, I weighed beetles before and after vein cutting, and simulated beetle feeding on leaves with and without vein cuts. By severing veins, the beetles decreased their total ingestion of exudate by at least 92%.     

Glucosinolates on the leaf surface perceived by insect herbivores: review of ambiguous results and new investigations

Herbivorous insects identify their host plants either by structural features, chemical cues, or a combination. Some insects probe the host leaf prior feeding or oviposition, other species use olfactorial cues or compounds somewhere on the surface. Insects attacking Brassicaceae are no exception, some are attracted and stimulated by volatile isothiocyanates (ITC), many others depend fully on the non-volatile glucosinolates (GS) for host-plant recognition and acceptance. Since most insects have no access to the leaf interior investigators concluded that GS must be present on the leaf surface and ITC in the headspace. However, peelings of mechanically removed surface waxes were devoid of measurable amounts of GS, whereas solvent surface extractions revealed a correlation between stomatal conditions and GS concentrations. Both observations lead to the conclusion that the presence of GS on the top leaf surface is rather unlikely. In the experimental part we show that a chloroform/methanol/water (2:1:1 vol/vol/vol) solvent leaf extract contains GS and, in addition, thia-triaza-fluorenes (TTF), other oviposition stimulants of the cabbage root fly, Delia radicum. Electrophysiological investigations showed that both, GS and TTF stimulated specific receptor neurones of the fly. We suggest that these compounds probably originated from deeper leaf layers and that herbivorous insects may penetrate the wax layer and perceive the stimulating compounds in deeper layers or through the stomata.     

Allometric analysis of the induced flavonols on the leaf surface of wild tobacco (Nicotiana attenuata)

Trichomes excrete secondary metabolites that may alter the chemical composition of the leaf surface, reducing damage caused by herbivores, pathogens and abiotic stresses. We examined the surface exudates produced by Nicotiana attenuata Torr. Ex Wats., a plant known to contain and secrete a number of secondary metabolites that are toxic or a deterrent to herbivorous insects. Extractions specific to the leaf surface, the trichomes, and the laminar components demonstrated the localization of particular compounds. Diterpene glycosides occurred exclusively in leaf mesophyll, whereas nicotine was found in both the trichomes and mesophyll. Neither rutin nor nicotine was found on the leaf surface. Quercetin and 7 methylated derivatives were found in the glandular trichomes and appeared to be excreted onto the leaf surface. We examined the elicitation of these flavonols on the leaf surface with a surface-area allometric analysis, which measures changes in metabolites independent of the effects of leaf expansion. The flavonols responded differently to wounding, methyl jasmonate (MeJA), herbivore attack and UV-C radiation, and the response patterns corresponded to their compound-specific allometries. Finding greater amounts of quercetin on younger leaves and reduced amounts after herbivore feeding and MeJA treatment, we hypothesized that quercetin may function as an attractant, helping the insects locate a preferred feeding site. Consistent with this hypothesis, mirids (Tupiocoris notatus) were found more often on mature leaves sprayed with quercetin at a concentration typical of young leaves than on unsupplemented mature leaves. The composition of metabolites on the leaf surface of N. attenuata changes throughout leaf development and in response to herbivore attack or environmental stress, and these changes are mediated in part by responses of the glandular trichomes.     

Sucking herbivore assemblage composition on greenhouse <Emphasis Type="Italic">Ficus</Emphasis> correlates with host plant leaf architecture

Understanding arthropod herbivore selectivity trends towards host plant attributes is essential for predicting plant-associated herbivore assemblage structure. Little is known about such interactions between spontaneous herbivore species and cultivated plants under specific conditions of botanical garden greenhouses. In this study, the taxonomic and functional composition of sucking arthropod herbivore assemblages were correlated with leaf anatomical and surface features of 33 host species of Ficus L. (Moraceae) across four distantly located greenhouse complexes of botanical gardens. The analyses revealed that the species number and abundance of scale insects and their individual families, sessile phloem feeders, total phloem feeders and total herbivores were significantly positively correlated with the thickness of leaf lamina, epidermis and mesophyll, stomatal length and width, and the presence of abaxial multiple epidermis and weak ferruginous non-glandular trichomes. Significant negative correlations were revealed between the same herbivore parameters and the density of glandular trichomes. Heliomorphic leaves supported higher abundance and species richness of sessile phloem feeders compared to sciomorphic ones. The parameters of some phloem and mesophyll feeder taxa also correlated with non-glandular trichome length and density, type of trichomes and epicuticular wax layer, and the presence of calcium oxalate crystals in the epidermis. Results of the study suggest that the leaf architecture-related herbivory trends under greenhouse conditions are similar to those occurring in natural ecosystems when considering the functional significance of particular leaf traits, and remain relevant at the scale of particular plant taxa with disregard of spatial factor.     

Changes in the amino acid composition and protein modification in phloem sap of rice

Pure phloem sap was collected from insects feeding on rice (Oryza sativa L.) leaves by a laser technique similar to the aphid stylet technique. Rapid circulation of nitrogen in the sieve tubes was demonstrated directly using ¹⁵N as a tracer. Application to the roots of the metabolic inhibitors of amino acids, aminooxyacetate and methioninesulfoximine, changed the amino acid composition in the sieve tubes. Feeding methionine to leaf tips resulted in its bulk transfer into the sieve tubes. In vitro experiments confirmed the existence of protein kinases in the pure rice phloem sap. The phosphorylation status of the sieve tube sap proteins was affected by the light regime. The possibility that changes in chemical composition or protein modification such as phosphorylation in the sieve tubes might affect plant growth are discussed.Analysis of pure phloem sap collected from rice plants by insect laser technique has shown dynamic changes in the chemical composition and the quality of proteins in the sap.     

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。