Immunohistochemical evidence for multiple photosystems in box jellyfish

Cubomedusae (box jellyfish) possess a remarkable visual system with 24 eyes distributed in four sensory structures termed rhopalia. Each rhopalium is equipped with six eyes: two pairs of pigment cup eyes and two unpaired lens eyes. Each eye type probably captures specific features of the visual environment. To investigate whether multiple types of photoreceptor cells are present in the rhopalium, and whether the different eye types possess different types of photoreceptors, we have used immunohistochemistry with a range of vertebrate opsin antibodies to label the photoreceptors, and electroretinograms (ERG) to determine their spectral sensitivity. All photoreceptor cells of the two lens eyes of the box jellyfish Tripedalia cystophora and Carybdea marsupialis displayed immunoreactivity for an antibody directed against the zebrafish ultraviolet (UV) opsin, but not against any of eight other rhodopsin or cone opsin antibodies tested. In neither of the two species were the pigment cup eyes immunoreactive for any of the opsin antibodies. ERG analysis of the Carybdea lower lens eyes demonstrated a single spectral sensitivity maximum at 485 nm suggesting the presence of a single opsin type. Our data demonstrate that the lens eyes of box jellyfish utilize a single opsin and are thus color-blind, and that there is probably a different photopigment in the pigment cup eyes. The results support our hypothesis that the lens eyes and the pigment cup eyes of box jellyfish are involved in different and specific visual tasks.     

Evolving eyes

Despite the incredible diversity among extant eyes, laws of physics constrain how light can be collected resulting in only eight known optical systems in animal eyes. Surprisingly, all animal eyes share a common molecular strategy using opsin for catching photons, but there are a diverse collection of mechanisms with proteins unrelated to each other used to focus light for vision. However, opsin is expressed in either one of two types of photoreceptor that differ fundamentally in their structure and tissue of origin. Taken together, this collection of observations strongly suggests that eyes have had multiple origins with remarkable convergence due to physics and molecular conservation of the opsin protein. Yet recent work has shown that a family of conserved genes are involved in eye formation despite substantial differences in their structure and origin, leading to a controversy over whether eyes evolved once or repeatedly. A likely resolution of this discussion is that particular genes and genetic programs have become associated with specific features needed for eyes and such suites of genes have been recruited as new eyes evolve. Since specific genes and their products are used repeatedly, it is somewhat difficult to conceptualize their causal relationships relative to evolutionary processes. However, detailed comparison of developmental programs may offer clues about multiple origins.     

Flight Distance and Eye Size in Birds

Larger eyes capture more information from the environment than small eyes, but also require more brain space for information processing. Therefore, individuals have to optimize the size of their eyes, leading to the prediction that larger eyes should have evolved in species with greater benefits from large eyes, such as species subject to intense predation risk. In a comparative analysis of 97 bird species, we found that species that fled at longer distances from an approaching potential predator indeed had relatively large eyes for their body size. In contrast, there was no indication that large eyes had evolved in species living in secluded habitats, or in species eating mobile prey. These findings are consistent with the assumption that eye size is labile and can evolve in response to changing predator environments. They also suggest that eye size may act as a constraint on optimal anti‐predator behavior, if the predator community changes as a consequence of introductions or invasions.     

The lens eyes of the box jellyfish <Emphasis Type="Italic">Tripedalia cystophora</Emphasis> and <Emphasis Type="Italic">Chiropsalmus sp.</Emphasis> are slow and color-blind

Box jellyfish, or cubomedusae, possess an impressive total of 24 eyes of four morphologically different types. Compared to other cnidarians they also have an elaborate behavioral repertoire, which for a large part seems to be visually guided. Two of the four types of cubomedusean eyes, called the upper and the lower lens eye, are camera type eyes with spherical fish-like lenses. Here we explore the electroretinograms of the lens eyes of the Caribbean species, Tripedalia cystophora, and the Australian species, Chiropsalmus sp. using suction electrodes. We show that the photoreceptors of the lens eyes of both species have dynamic ranges of about 3 log units and slow responses. The spectral sensitivity curves for all eyes peak in the blue-green region, but the lower lens eye of T. cystophora has a small additional peak in the near UV range. All spectral sensitivity curves agree well with the theoretical absorbance curve of a single opsin, strongly suggesting color-blind vision in box jellyfish with a single receptor type. A single opsin is supported by selective adaptation experiments.     

Ultrastructure of pigmented adult eyes in errant polychaetes (Annelida): implications for annelid evolution

The evolution of photoreceptor cells and eyes in Metazoa is far from being resolved, although recent developmental and structural studies have provided strong evidence for a common origin of photoreceptor cells and existence of sister cell types already in early metazoans. These sister cell types are ciliary and rhabdomeric photoreceptor cells, depending on which part of each cell is involved in photoreception proper. However, a crucial point in eye evolution is how the enormous structural diversity of photoreceptor cells and visual systems developed, given the general molecular conservation of the photoreceptor cells. One example of this diversity can be observed in Annelida. Within the polychaetes the errant forms, taxon Aciculata, constitute the only group possessing true multicellular eyes in the adult stage. Thus far these organs have been investigated only in taxa of Phyllodocida, a subgroup of Aciculata. Data on Eunicida and Amphinomida as well as certain phyllodocidan taxa had been lacking. The ultrastructure of these adult eyes was investigated in various species of errant polychaetes, belonging to Amphinomidae, Eunicidae and Hesionidae, to elucidate whether they provide any phylogenetic clues regarding either the evolution of visual systems in Annelida or lophotrochozoan phylogeny in general. These eyes are composed of numerous supportive pigment cells and rhabdomeric photoreceptor cells and sometimes additional cell types. As a rule the pigment and rhabdomeric cell types form a continuous epithelium in which the two types intermingle. Presence of granules with shading pigment in sensory cells is a common feature but is apparently restricted to a taxon comprising Phyllodocida and Eunicida s. str. Very likely a lens-like structure does not belong to the ground pattern of annelid eyes, despite its presence in Phyllodocida. These lens-like structures are formed by secretions or cellular processes of the pigment cells. In many species the eye cup communicates with the exterior via a small cuticularized canal. This canal is interpreted as a rudiment due to the mode of formation in the epidermis. With respect to current phylogenetic hypotheses, these multicellular eyes have either been developed in the stem species of a taxon Aciculata nested within the polychaetes or have been evolved in the stem lineage of Annelida. Similarities to gastropod eyes are interpreted as convergent and not as indication of common origin. Except for the photoreceptor cells proper, the structure of the adult eyes in polychaetes most likely does not help to resolve lophotrochozoan phylogeny.     

DO IMAGE-FORMING EYES PROMOTE EVOLUTIONARY DIVERSIFICATION?

It has been suggested that image-forming eyes promote the evolutionary diversification (measured by species richness) of the groups that possess them. Several different processes could give rise to this effect, including diversifying selection in a new adaptive zone (or zones) and a reduced rate of extinction due to enhanced competitive abilities. I tested the generality of the hypothesis that imaging eyes increase net speciation by comparing extant species numbers of 12 groups that have such eyes (as categorized by Land and Fernald 1992) with those of their cladistic sister groups that lack such organs. Even assuming the published hypotheses of phylogenetic relationships that most favor increased net speciation of visual groups, these comparisons show no significant association between imaging eyes and species richness. Increased activity, as indicated by published accounts of locomotory speed, is significantly associated with the evolution of image-forming eyes. This suggests that a large “visual adaptive zone” might be characterized by relatively high activity. However, when diversity comparisons are limited to eight cases in which the evolution of imaging eyes is associated with increased activity, there is still no significant association between such eyes and species richness. The fossil record indicates that the only visual groups that have undergone major evolutionary radiations evolved imaging eyes early in the history of metazoans (before the Silurian). The radiations of these early groups may have largely filled up niches for visual animals and thus prevented the subsequent proliferation of other groups with image-forming eyes. Alternatively, it may be that image-forming eyes have no exceptional effect on diversification or that their effects are obscured by other factors in the long run.     

Sense organs in Pycnogonida: A review

The Pycnogonida or sea spiders are exclusively marine invertebrates, numbering about 1,300 described species worldwide. Given their remarkable position in phylogeny as basal chelicerates or even basal euarthropods, the structure of their sense organs can reveal important characters, which—in a comparative framework—provide arguments to phylogenetic discussions and help to develop scenarios of evolutionary transformations. This review summarizes current knowledge and presents new original data on the sense organs in pycnogonids, that is, the eyes, the lateral sense organs and the ciliary or sensillar sense organs. Except for the eyes, there are not many detailed studies available. The ultrastructure of the R‐cells of the four eyes located on the ocular tubercle is described as “pseudoinverted”. The eyes are innervated to two visual neuropils located in the protocerebrum. The features of the lateral sense organ, also located on the ocular tubercle, are hitherto not conclusively resolved, a chemo‐ or thermoreceptive function is suggested. Finally, an overview of the various ciliary or sensillar sense organs distributed all over the body is given and the fine structure of branched setae is shown for the first time. The morphology of the sense organs of pycnogonids is compared with that of other arthropod taxa and assessed against the background of current theories of arthropod evolution.     

Visual fields and eye morphology support color vision in a color-changing crab-spider

Vision plays a major role in many spiders, being involved in prey hunting, orientation or substrate choice, among others. In Misumena vatia, which experiences morphological color changes, vision has been reported to be involved in substrate color matching. Electrophysiological evidence reveals that at least two types of photoreceptors are present in this species, but these data are not backed up by morphological evidence. This work analyzes the functional structure of the eyes of this spider and relates it to its color-changing abilities. A broad superposition of the visual field of the different eyes was observed, even between binocular regions of principal and secondary eyes. The frontal space is simultaneously analyzed by four eyes. This superposition supports the integration of the visual information provided by the different eye types. The mobile retina of the principal eyes of this spider is organized in three layers of three different types of rhabdoms. The third and deepest layer is composed by just one large rhabdom surrounded by dark screening pigments that limit the light entry. The three pairs of secondary eyes have all a single layer of rhabdoms. Our findings provide strong support for an involvement of the visual system in color matching in this spider.     

Secondary eyes mediate the response to looming objects in jumping spiders (Phidippus audax,Salticidae)

Some species have sensory systems divided into subsystems with morphologically different sense organs that acquire different types of information within the same modality. Jumping spiders (family Salticidae) have eight eyes. Four eyes are directed anteriorly to view objects in front of the spider: a pair of principal eyes track targets with their movable retinae, while the immobile anterior lateral (AL) eyes have a larger field of view and lower resolution. To test whether the principal eyes, the AL eyes, or both together mediate the response to looming stimuli, we presented spiders with a video of a solid black circle that rapidly expanded (loomed) or contracted (receded). Control spiders and spiders with their principal eyes masked were significantly more likely to back away from the looming stimulus than were spiders with their AL eyes masked. Almost no individuals backed away from the receding stimulus. Our results show that the AL eyes alone mediate the loom response to objects anterior to the spider.     

珍珠菜属植物种皮微形态特征及其系统学意义

利用光学显微镜和扫描电镜对11种珍珠菜属植物种皮微形态特征进行了研究。根据种皮微形态特征,可以明显的划分为两种类型,即黑腺珍珠菜型和金爪儿型。珍珠菜亚属的7种植物均属于黑腺珍珠菜型,黄连花亚属的4种植物均属于金爪儿型。种皮微形态特征支持以雄蕊群的构造为主要依据的属下分类系统,在亚属级水平具有重要的分类学意义。     

The optical sensitivity of compound eyes: theory and experiment compared

The Land sensitivity equation is a well-known tool for comparing optical performance between eyes. Despite this, the equation has never been experimentally tested. Here, we present, to our knowledge, the first experimental validation of the equation. We have investigated different insect species active at different intensities, and possessing different types of compound eyes, to compare ratios of calculated sensitivities to ratios determined experimentally. Experimental optical sensitivities were measured by adjusting the intensity of an external light source until photoreceptors in the different eyes produced roughly equal numbers of photon responses ('bumps') per second. The sensitivity ratios obtained in this manner agree well with those obtained using the equation. We conclude that the Land equation remains an excellent tool for comparing sensitivities between different eyes.     

The frontal eyes of crustaceans

Frontal eyes of crustaceans (previously called nauplius eye and frontal organs) are usually simple eyes that send their axons to a medial brain centre in the anterior margin of the protocerebrum. Investigations of a large number of recent species within all major groups of the Crustacea have disclosed four kinds of frontal eyes correlated with taxonomic groups and named after them as the malacostracan, ostracod-maxillopodan, anostracan, and phyllopodan frontal eyes. The different kinds of eyes have been established using the homology concept coined by Owen [Owen, R., 1843. Lectures on the comparative anatomy and physiology of the invertebrate animals. Longman, Brown, Green, Longmans, London] and the criteria for homology recommended by Remane [Remane, A., 1956. Die Grundlagen des natürlichen Systems, der vergleichenden Anatomie und der Phylogenetik. 2nd ed. Akademische Verlagsgesellschaft, Geest und Portig, Leipzig]. Common descent is not used as a homology criterion. Frontal eyes bear no resemblance to compound eyes and in the absence of compound eyes, as in the ostracod-maxillopodan group, frontal eyes develop into complicated mirror, lens-mirror, and scanning eyes. Developmental studies demonstrate widely different ways to produce frontal eyes in phyllopods and malacostracans. As a result of the studies of recent frontal eyes in crustaceans, it is concluded by extrapolation that in crustacean ancestors four non-homologous frontal eye types evolved that have remained functional in spite of concurrent compound eyes.     

Novel organelle associations in photoreceptors of a serpulid polychaete worm

Cerebral and branchial eyes of serpulid polychaetes have been studied by electron microscopy for the first time. In one species both eye types possess a novel ultrastructure. The receptor cells of the simple cerebral ocelli are rhabdomeric and display a close structural and functional relationship between ciliary rootlets and mitochondria. The receptors of the compound branchial eyes contain both a rhabdom and a stack of photosensitive ciliary membranes.     

Zur genetischen interpretation morphologischer gesetzmäßigkeiten der degenerativen evolution

Complex organs such as the eyes of vertebrates are based upon highly polygenic systems. So, remarkable differences in the eyes of cave fishes and their overground ancestors are caused by several mutations. Although in cavernicolous populations natural selection seems to play no part in eye degeneration, there is evidence for morphologic regularities in the degenerative process. These morphologic regularities are due to pleiotropic effects of the mutating genes.     

Impact of habitat and life trait on character evolution of pallial eyes in Pectinidae (Mollusca: bivalvia)

Pectinidae, a large group of marine bivalves comprising more than 300 species worldwide, inhabit a diverse array of habitats, enabling an enormous radiation, and yielding many different life forms and adaptations. This apparent diversity led to the distinction of ecotypes based on shell morphology and lifestyle. Eyes in Pectinidae (Bivalvia, Pteriomorphia) have long sparked scientific interest and have been described for various species over the past two centuries. These eyes are morphologically and functionally highly complex. Despite this complexity, studies have focused mostly on functional aspects with only few examining the relationships associated with different environmental or evolutionary traits. Here, the pallial eye structure within the Pectinidae was examined using Masson Goldner Trichrom staining, and ancestral character estimation with BayesTraits was performed to reconstruct macro-evolutionary patterns. To evaluate the connection of substrate type and lifestyle to the evolution of eyes, we compared eyes within the major subgroups of Pectinidae while considering the different lifestyles and substrate types as well as different depth ranges. The results indicate a tendency towards a taxon-/clade-specific evolution in respect to characters such as the cornea and lens while depth specific adaptations occur mainly in the light sensitive compartments of the retina. Successive reduction of eyes seems to occur from shallow to deep water species and ends in a total reduction of all structures in deep sea species.     

青海孟达国家级自然保护区植物多样性研究

对孟达国家级自然保护区的植物多样性进行了研究。结果表明:保护区内有植物90科、298属、529种(包括变种)。其中,有裸子植物16种,被子植物488种,蕨类植物17种,苔藓类植物8种。国家珍稀濒危保护植物8种。资源植物丰富,分为药用、观赏、食用等10大类。植物垂直分布明显,生态系统类型多样。提出保护植物多样性应采取的措施。     

Deschampsia antarctica: genetic and molecular-biological aspects of spreading in Antarctica

The genetical and molecular biological attributes of Deschampsia antarctica are considered in relation to its remarkable success in the colonization of the maritime Antarctic. However, none of the data from several published research studies provide convincing evidence that this species possess any obvious unique or specialised adaptations which might account for this success. While its occurrence appears to have spanned much of the Holocene, there is no evidence that there have ever been any other native vascular plant species in the maritime Antarctic. It is still unclear why no other taxa have succeeded in colonising this region. Further intensive studies of genetics and molecular biology of this species, and of other potential immigrants, may provide a better understanding of their enigmatic success in the maritime Antarctic.     

Game theory provides no explanation for seed size variation in grasslands

Game theoretical models have been suggested to explain the maintenance of a remarkable variation in seed size across species in most types of vegetation. According to these models, which are based on the existence of a trade-off between seed size and seed number; smaller-seeded species can invade any species mixture due to their numerical advantage, and larger-seeded species can invade any species mixture due to their competitive superiority over smaller-seeded species during recruitment. However, till now, there is very limited evidence for seed size effects on recruitment interactions among different species. An experiment was designed using 16 species in Swedish grasslands, varying 384-fold in seed size. Species were sown pairwise and alone in disturbed versus undisturbed small plots in grazed versus ungrazed grassland, and the resulting recruitment was recorded. Seedling densities in the experimental plots were within the natural range. Both disturbance and grazing had a positive effect on recruitment. Seed size did not affect recruitment except from a tendency that increasing seed size made recruitment less dependent on disturbance. Recruitment of smaller-seeded species was not affected by the presence of larger-seeded species. Larger-seeded species did not generally win in direct contest. These results suggest that game theoretical models do not explain maintenance of seed size variation across species in these grasslands. Alternative explanations for seed size variation are that either small-scale heterogeneity provide conditions favouring a range of different seed sizes or other attributes than seed size effectively determine recruitment.