黑带食蚜蝇Episyrphus balteatus De Geer的复眼结构及其调光机制

【目的】观察研究黑带食蚜蝇Episyrphus balteatus De Geer成虫复眼形态、小眼结构和不同光暗条件对小眼结构的影响,以明确其光视觉的结构基础和调光机制。【方法】利用组织切片法和扫描电镜等技术。【结果】1.复眼位于头部两侧,正面观呈半球形,占据除额颜外大部分头部。雄虫与雌虫单个复眼分别有约7 180个、7 230个小眼。各小眼面呈整齐排列的规则六边形。2.小眼由角膜及伪晶锥组成的屈光器、不同水平面分布的8个小网膜细胞及其特化形成的离散型视杆、屏蔽色素细胞和基膜等组成。小眼自远端至近端由主色素细胞和12个附属色素细胞围绕。3.随光暗条件的改变小眼内的附属色素细胞色素和基细胞细胞核沿小眼纵轴移动。光适应时,附属色素细胞色素颗粒沿小眼纵轴均匀分布,基细胞细胞核位于基膜上方。暗适应时,附属色素细胞色素颗粒向伪晶锥近端压缩,基细胞细胞核亦向远端移动,到达视杆中段。【结论】黑带食蚜蝇复眼精密的小眼排列形式和内部结构均显示了其强大的生理功能;屏蔽色素颗粒的移动是其复眼适应外界光环境变化的重要机制。本试验为进一步探究黑带食蚜蝇视觉结构和光调节机制,以及与其飞行行为间的关系提供了一定的理论基础。     

黑翅土白蚁有翅成虫复眼的形态结构

采用组织切片法光镜下观察黑翅土白蚁Odontotermes formosanus(Shiraki)有翅成虫的复眼形态结构及光、暗适应条件下色素颗粒移动的规律。结果如下:(1)头正前方观,复眼外部形态略呈圆形。(2)有翅成虫复眼类型属于并列像眼,每只复眼约由360个小眼组成。(3)每个小眼是由1套屈光器(1个角膜和1个晶锥)、小网膜色素细胞、视杆和基细胞等几部分组成。小网膜色素细胞内均含有丰富的色素颗粒。(4)在光适应条件状态下,屈光器及视杆周围的色素颗粒主要分布在视杆部位的上侧,暗度适应条件状态时则较均匀地分布于视杆两侧上下;性别对色素颗粒分布无明显影响。     

夜蛾趋光特性的研究——向灯飞原因的进一步分析

在灯区内亮灯之后,棉铃虫蛾和粘虫蛾的复眼状况受照射灯光的影响。 一、灯距与复眼色素分布有以下关系: 二.在100—5米范围内,每当灯距减小一半,直射光强增强到原来的四倍时,复眼色素变化只有原来的1—2.36倍。 2.在上述范围内,直射光强的变幅要比虫眼所体现的相应变幅大200倍左右,表明虫眼对光强变化有相当大的调节和适应能力。 二、受光方向不同,能使同一个体的两眼处于不同状态。 三、上灯的棉铃虫、粘虫、小地老虎和毒蛾,同一个体的两眼状态基本一致,说明接受光照条件相同;复眼状况从形态上看介于充分光、暗适应之间,而偏近于暗适应状况。 根据光照与复眼状况的相关性,对夜娥类的扑灯原因和灯区的有效范围,进行了初步探讨。     

夜蛾趋光特性的研究:复眼转化过程中的行为变异

影响蛾类趋光性和灯光诱蛾效果的因素很多。过去,国内外许多观察和实验都侧重在研究光源的辐射光谱和辐射强度以及气候条件的影响。至于蛾类的主要感光器官的适应状态怎样影响趋光性,却极少引起注意。 陈元光等(1963)曾经用电生理学的方法,测出粘虫蛾复眼的暗适应过程大约为30—45分钟。Agee(1972)用电生理方法研究复眼,发现美国棉铃虫复眼在暗适应中,达到最高灵敏度的时间为30分至285分,雌、雄的平均时间分别为88分和115分;美国烟青虫雌、雄蛾的平均时间分别为119分和121分。最近,李典谟等(1977)以复眼反射光的增强作为暗适应的指标,测出烟青虫蛾在暗室内暗适应时,复眼的转化速率很不一致。快的个体约经过1小时就达到最大反射值;慢的个体则需经过3小时,而绝大多数个体(79％)则在1—2(1/2)小时内。因此,他们推测在入夜后的一段时间里,田间昆虫的眼由昼眼转化为夜眼的情况是各不相同的,并且认为这种差异会影响成虫对于灯光发生不同的行为反应。棉铃虫(Heliothis armigera)蛾是灯光防治的主要对象之一,为了进一步探索这个问题,我们认为有必要检查一下在日夜交替的自然暗适应条件下其复眼状态究竟是否一致,复眼状态怎样影响着趋光性,以便深入地了解夜蛾类趋光性的控制因素。     

棉铃虫和玉米螟成虫复眼光反应特性的比较研究

通过记录棉铃虫和玉米螟复眼的网膜电图（Ｅｌｅｃｔｒｏｒｅｔｉｎｏｇｒａｍ,ＥＲＧ）比较了二者对光刺激的反应特性。结果如下：（１）二者具有相似的光谱反应曲线,均对５６２ｎｍ的光最敏感,另外对４００ｎｍ、４８３ｎｍ光也较敏感;（２）二者对这三种敏感光的光强度曲线近似,在一定范围内均近似线性递增,但玉米螟复眼ＥＲＧ在５６２ｎｍ和４８３ｎｍ下具有更高的反应幅度;（３）二者对白光的光强度曲线也近似,但随复眼适应状态及测试时间的不同,表现出不同的光敏感性。在相同条件下,棉铃虫复眼表现出更强的敏感性;同种昆虫复眼夜间比白天敏感,暗眼比亮眼敏感;（４）对不同测试时间复眼明暗状态转化的比较表明,棉铃虫复眼的ＥＲＧ反应具有更强的昼夜节律性。     

螺旋粉虱成虫的复眼形态及其内部结构

采用扫描电镜和组织切片法,观察了螺旋粉虱Aleurodicus dispersus Russell成虫复眼的形态及其显微结构。结果表明,螺旋粉虱复眼半球状,呈“∞”形分布于头部两侧,单个复眼约由253个小眼组成;各小眼面微凸,复眼中心区域小眼多为规则的六边形,密集排列似蜂窝状;近背区边缘小眼多为五边形或近圆形,小眼排列疏松,且少量相邻小眼的间距较大。雌、雄复眼小眼面积约为85μm2。单个小眼由角膜、晶体、网膜细胞及其特化产生的视杆和基细胞等几部分组成。晶体有四个晶锥细胞构成,晶体、视杆周围和色素细胞内均含有大量的色素颗粒。螺旋粉虱的复眼属于并置复眼。光、暗条件下,小眼的色素颗粒分布有所不同。光适应条件下,色素颗粒较均匀地分布于视杆上下两侧;暗适应状态下,色素颗粒则主要分布在视杆上侧和晶体下侧。而在相同的明、暗适应条件下,性别对色素颗粒的分布无显著影响。     

蝗虫复眼光谱敏感性的比较研究——Ⅰ.屏蔽色素显微分光光谱的测量

用显微分光光谱法,测定了9种蝗虫复眼两类屏蔽色素的光密度。咖啡色屏蔽色素在400—520nm 波段密度最高,在深红区密度最低。反光屏蔽色素在365—390nm 波段的密度最高,在深红区最低。此外还测定了9种蝗虫之一的黄脊蝗复眼的光屏蔽色素的反射光谱,结果显示在近紫外和橙红区反光率最高。关于屏蔽色素对蝗虫视觉功能如光谱敏感性、明适应、视锐度、视色素和间视紫红之间的转化等问题曾加以讨论。     

环境照度和温度对两种夜蛾复眼适应转化的影响

粘虫蛾复眼经过六种不同照度(0.5、1.5、6.0、13、29、50勒克司)的适应所测得的各反光区指数表明,当环境的照度增加时,大多数蛾眼受到光的影响而反光区的面积明显变小。在29勒克司的照度环境时,则大部分因受到光的抑制而不出现反光区。当低于29勒克司时,复眼的适应状态就参差不齐,这可能反映了粘虫蛾的复眼并不是都具备同等适应的能力。在50勒克司的照度下,蛾眼全都被抑制而停留在昼眼状态。 棉铃虫蛾复眼在七种温度(1、5、8、9、10、15、25℃)下经过完全暗适应的处理后表明,随着温度的提高,蛾眼的平均反光区面积也相应增大。低于8℃以下的温度时,对蛾眼转化有明显差异,在8℃的温度下暗适应1.5小时后,大多数的蛾眼正处在转化或未转化的状态,因此可以认为,在8℃时棉铃虫蛾复眼的视觉灵敏度不是很高的。同样大小的反光区面积,当温度低时通过蛾眼底片的相对透射光量就大,温度高时通过的相对透射光量就较少。     

龟纹瓢虫成虫的复眼形态及其显微结构

利用光镜、组织切片法观察了龟纹瓢虫Propylaea japonica(Thunberg)成虫的复眼形态及其显微结构。结果如下:(1)头正前方观,复眼外形似半球,且后方稍向内合拢。每个复眼约包括630个小眼。(2)每个小眼是由1套屈光器(1个角膜和1个晶锥)、6至8个小网膜细胞及其特化产生的视杆和基细胞等几部分组成。晶体周围及小网膜色素细胞内均含有丰富的色素颗粒。(3)小眼整体纵切显示,其上、下段色素颗粒分布相对较多,中段分布较少。(4)明、暗适应状态对小眼的色素颗粒分布有影响,性别对其分布无明显影响。明适应状态下,其色素颗粒较均匀地分布于视杆两侧上下,暗适应状态时色素颗粒则主要分布在视杆部位的上侧,显示其具有一定的重叠眼性质;而在相同的明、暗适应状态下其雌、雄成虫复眼的色素颗粒分布间无明显差异。     

异色瓢虫显现变种复眼的形态、显微结构及其光暗条件下的适应性变化

复眼是昆虫的主要视觉器官,对于其寻找食物、配偶、栖息场所以及学习记忆等活动具有重要作用。本研究采用扫描电镜和石蜡切片技术对异色瓢虫显现变种Harmonia axyridis ab. conspicua复眼的外部形态和内部显微结构进行了观察。结果发现：(1)复眼近椭圆形,位于头部两侧,触角窝处有缺刻,小眼表面光滑平坦,无角膜乳突结构。其雌、雄成虫复眼的小眼数分别约为705和691;(2)复眼中心区域小眼呈六边形,排列紧密,边缘区域的小眼为不规则的四边形或五边形; (3)每个小眼由角膜、晶锥、8个小网膜细胞、视杆、基膜以及色素细胞组成。晶锥由4个晶锥细胞构成,8个小网膜细胞中6个位于边缘、2个位于中央;(4)暗条件下复眼显微结构存在明显差异：光适应条件下,色素颗粒主要分布在晶锥和视杆交界处的周围,周围视杆呈环形,内、外两侧均被色素颗粒包围;暗适应条件下,色素颗粒发生纵向移动,均匀地分布在晶锥和视杆的周围,周围视杆发生扭曲呈不规则的多角形,仅外侧有色素颗粒分布。结果表明,异色瓢虫显现变种的复眼属于并列复眼,可通过色素颗粒的纵向移动以及周围视杆扭曲变形等机制来适应外界明暗环境的变化。     

亚洲玉米螟蛾趋光行为及复眼结构节律性研究

用行为实验和光学显微镜观察比较了夜间暗适应处理和日间暗适应处理的亚洲玉米螟蛾Ostrinia furnacalis(Guenee)趋光反应及小眼的显微结构。(1)夜间暗适应处理组的趋光反应率明显高于日间暗适应处理组。5个照度行为实验中,前者最高反应率达80%,最低23%,总反应率近48%,而后者分别只有35%、10%和21%。(2)夜间暗适应与日间暗适应的复眼其屏蔽色素分布大体相同,但小网膜细胞核的分布区域有明显差异,表明亚洲玉米螟蛾小眼结构变化存在一定的节律性。这种结构节律变化可能是导致趋光反应昼夜差异的原因之一。     

Diurnal changes in angular sensitivity of crab photoreceptors

Summary The electrophysiological and anatomical consequences of diurnal changes in screening pigment position were investigated in the apposition eye of the portunid crabScylla serrata. Intracellular recordings revealed that the acceptance angles of dark-adapted photoreceptors enlarged up to four-fold at night compared with photoreceptors dark-adapted in the day. Furthermore, while light adaptation at night caused acceptance angles to narrow, dark adaptation in the day caused no significant broadening of angles. These electrophysiological changes correlated with pigment movements in the eye observed both histologically and in the deep pseudopupil. It is found that the distal pigment cells change diurnally so that the field-stop which these cells form in front of the photoreceptors is opened in the night and closed in the day time.One feature of the diurnal rhythm is that it prevents photoreceptor fields of view enlarging when eyes are dark adapted in the day. InScylla, photoreceptor fields of view take tens of minutes to narrow upon exposure of crabs to light at night. By preventing a similar broadening in the day, the diurnal rhythm may enable animals suddenly leaving dark refuges to be pre-adapted to daylight. To a range of species which utilise refuges such a mechanism would be of significant advantage, especially after disturbance by predators.We are grateful to Prof. G.A. Horridge for constant encouragement and to Drs. S.B. Laughlin, M. Wilson, S. Shaw and M.F. Land for helpful advice.     

飞蝗复眼生理和结构上的节律变化

采用细胞内记录和光镜方法研究了飞蝗(Locusta migratoria)夜间和日间在暗适应和明适应状态下小网膜细胞角敏感度以及晶锥和小网膜细胞之间区域结构上的变化.结果表明小网膜细胞角敏感度的变化不仅仅由于晶锥周围主色素细胞色素颗粒的移动,而且也由于小眼感杆束结构上的节律变化.     

锯缘青蟹视网膜电图特性的昼夜节律变化

本文系统地分析了锯缘青蟹scylla serrata视网膜电图特性(敏感度、光谱敏感性和波形)的昼夜节律性变化。这种节律性可能主要是由于视网膜屏蔽色素位置的昼夜变化所致,但也可能存在另一种机制。     

THE DARK ADAPTATION OF THE EYE OF THE HONEY BEE

Bees which are held in a fixed position so that only head movements can be made, respond to a moving stripe system in their visual field by a characteristic motion of the antennae. This reflex can be used to measure the bee''s state of photic adaptation. A curve describing the course of dark adaptation is obtained, which shows that the sensitivity of the light adapted bee''s eye increases rapidly during the first few minutes in darkness, then more slowly until it reaches a maximum level after 25 to 30 minutes. The total increase in sensitivity is about 1000 fold. The adaptive range of the human eye is about 10 times greater than for the bee''s eye. The range covered by the bee''s eye corresponds closely to the adapting range which is covered by the rods of the human eye.     

Studies on the Relation between the Pigment Migration and the Sensitivity Changes during Dark Adaptation in Diurnal and Nocturnal Lepidoptera

The functional significance of the pigment migration in the compound insect eye during dark adaptation has been studied in diurnal and nocturnal Lepidoptera. Measurements of the photomechanical changes were made on sections of eyes which had been dark-adapted for varying periods of time. In some experiments the sensitivity changes during dark adaptation were first determined before the eye was placed in the fixation solution. No change in the position of the retinal pigment occurred in Cerapteryx graminis until the eye had been dark-adapted for about 5 minutes. The start of the migration was accompanied by the appearance of a break in the dark adaptation curve. During longer periods of dark adaptation the outward movement of the pigment proceeded in parallel with the change in sensitivity, the migration as well as the adaptive process being completed within about 30 minutes. In the diurnal insects chosen for the present study (Erebia, Argynnis) the positional changes of the retinal pigment were insignificant in comparison with the movement of the distal pigment in Cerapteryx graminis. On the basis of these observations the tentative hypothesis is put forward that the second phase of adaptive change in nocturnal Lepidoptera is mediated by the migration of the retinal pigment while the first phase is assumed to be produced by the resynthesis of some photochemical substance. In diurnal insects which have no appreciable pigment migration the biochemical events alone appear to be responsible for the increase in sensitivity during dark adaptation.     

Daily changes of structure,function and rhodopsin content in the compound eye of the crabHemigrapsus sanguineus

The compound eye of the crab hemigrapsus sanguineus undergoes daily changes in morphology as determined by light and electron microscopy, both in the quantity of chromophore substances studied by HPLC and in visual sensitivity as shown by electrophysiological techniques. 1. At a temperature of 20 degrees C, the rhabdom occupation ratio (ROR) of an ommatidial retinula was 11.6% (maximum) at midnight, 8.0 times larger than the minimum value at midday (1.4%). 2. Observations by freeze-fracture revealed that the densities of intra-membranous particles (9-11 nm in diameter) of rhabdomeric membrane were ca. 2000/microns 2 and ca. 3000/microns 2 for night and daytime compound eyes, respectively. 3. Screening pigment granules migrated longitudinally and aggregated at night, but dispersed during the day. Reflecting pigment granules migrate transversally in the proximal half of the reticula layer i.e. cytoplasmic extensions containing reflecting pigment granules squeeze between neighbouring retinula cells causing optical isolation (Fig. 4). Thus the screening pigment granules within the retinula cells show longitudinal migration and radial movement so that the daytime rhabdoms are closely surrounded by the pigment granules. 4. At 20 degrees C, the total amount of chromophore of the visual pigment (11-cis and all-trans-retinal) was 1.4 times larger at night than during the day i.e. 46.6 pmol/eye at midnight and 33.2 pmol/eye at midday. Calculations of the total surface area of rhabdomeric membrane, total number of intra-membranous particles in rhabdomeric membrane and the total number of chromophore molecules in a compound eye, indicate that a considerable amount of chromophore-protein complex exists outside the rhabdom during the day. 5. The change in rhabdom size and quantity of chromophore were highly dependent on temperature. At 10 degrees C both rhabdom size and amount of chromophore stayed close to daytime levels throughout the 24 hours. 6. The intracellularly determined relative sensitivity of the dark adapted night eye to a point source of light was about twice as high as the dark-adapted day eye. Most of the increase in the sensitivity is attributed primarily to the effect of reflecting pigment migration around the basement membrane and, secondarily, to the changes in the amount and properties of the photoreceptive membrane. The results form the basis of a detailed discussion as to how an apposition eye can function possibly as a night-eye.     

THE REGENERATION OF VISUAL PURPLE IN THE LIVING ANIMAL

1. The accumulation of visual purple in the retina after bleaching by light has been studied in the intact eye of the frog. The data show that duration and intensity of light adaptation, which influence the course of human dark adaptation as measured in terms of visual threshold, have a similar influence on the course of visual purple regeneration. 2. At 25°C. frogs which have been light adapted to 1700 millilamberts and then placed in the dark, show an increase in visual purple concentration which begins immediately and continues for 70 minutes until a maximum concentration is attained. The increase, although beginning at once, is slow at first, then proceeds rapidly, and finally slows up towards the end. Frogs which have been adapted to 9500 millilamberts show essentially the same phenomenon except that the initial slow period is strongly delayed so that almost no visual purple is formed in the first 10 minutes. 3. At 15°C. the initial delay in visual purple regeneration occurs following light adaptation to both 1700 and 9500 millilamberts. The delay is about 10 minutes and is slightly longer following the higher light adaptation. 4. The entire course of visual purple accumulation in the dark takes longer at the lower temperature than at the higher. The temperature coefficient for 10°C. is about 1.8. 5. In contrast to the behavior of the isolated retina which has small amounts of vitamin A and large amounts of retinene immediately after exposure to light, the intact eye has large amounts of vitamin A and little retinene after exposure to light for 10 minutes. In the intact eye during dark adaptation, the amount of vitamin A decreases markedly while retinene decreases only slightly in amount. If retinene is formed in the intact eye, the change from retinene to vitamin A must therefore occur rapidly in contrast to the slow change in the isolated retina. 6. The course of visual purple regeneration may be described by the equation for a first order autocatalyzed reaction. This supposes that the regeneration of visual purple is catalyzed by visual purple itself and accounts for the sigmoid shape of the data.     

Rhabdom size and photoreceptor membrane turnover in a muscoid fly

Summary The cross-sectional area of rhabdomeres in the compound eye of the blowfly, Lucilia, was found to remain constant under 12 h light/12 h dark cyclic lighting, and 10 days constant light or darkness. It was reduced only slightly during 3 h light after 10 days darkness (by 21%), or on exposure to 2h darkness + 1.5 h light after 10 days light (by 10%). Morphological evidence indicates that shedding of photoreceptor membrane during turnover is achieved by an extracellular route, and by pinocytosis from the bases of the microvilli. The photoreceptor membrane shed by both mechanisms appears to accumulate in multivesicular bodies. The amount of photoreceptor membrane shedding, as indicated by numbers of multivesicular bodies, is constant throughout the day and night on cyclic lighting, decreases in constant darkness, but returns to its normal level after an exposure to 3 h light subsequent to 10 days darkness.     

大草蛉成虫复眼的外部形态及其显微结构

用扫描电镜和光学显微镜观察了大草蛉Chrysopa pallens Ramber成虫复眼的外部形态及明、暗适应和性别对其显微结构的影响。结果发现：（1）其复眼呈半球形,位于头部两侧,略成“八”字形排列,单个复眼约由3 600个小眼组成,最前和最后小眼之间的夹角约为180°,最上和最下小眼之间的夹角约200°;（2）小眼主要由角膜、晶锥和6～8个小网膜细胞、基膜组成,外围环绕有2个初级虹膜色素细胞和6个次级虹膜色素细胞,基膜处有色素颗粒分布;（3）暗适应时,晶锥开裂程度较大,远端5～7个网膜细胞核向远端移动,与晶锥近端相接或接近,次级虹膜色素颗粒亦向远端移动包围晶锥;明适应时,晶锥开裂程度小或闭合,远端网膜细胞核向近端移动,透明带显现,大部分次级虹膜色素颗粒亦向近端移动分布在小网膜细胞柱周围,包被透明带;（4）在相同的明、暗适应下,雌、雄成虫复眼的显微结构无明显差异。结果表明大草蛉复眼为透明带明显的重叠象眼,其小眼不但具有次级虹膜色素颗粒纵向移动的常规调光机制,还存在晶锥开闭、远端网膜细胞核移动和基膜色素颗粒纵向扩散的调光新机制。