2016-11-02T18:49:00

Spiders are among the world's most species-rich animal lineages, and their visual systems are likewise highly diverse. These modular visual systems, composed of four pairs of image-forming "camera" eyes, have taken on a huge variety of forms, exhibiting variation in eye size, eye placement, image resolution, and field of view, as well as sensitivity to color, polarization, light levels, and motion cues. However, despite this conspicuous diversity, our understanding of the genetic underpinnings of these visual systems remains shallow. Here, we review the current literature, analyze publicly available transcriptomic data, and discuss hypotheses about the origins and development of spider eyes. Our efforts highlight that there are many new things to discover from spider eyes, and yet these opportunities are set against a backdrop of deep homology with other arthropod lineages. For example, many (but not all) of the genes that appear important for early eye development in spiders are familiar players known from the developmental networks of other model systems (e.g., Drosophila). Similarly, our analyses of opsins and related phototransduction genes suggest that spider photoreceptors employ many of the same genes and molecular mechanisms known from other arthropods, with a hypothesized ancestral spider set of four visual and four nonvisual opsins. This deep homology provides a number of useful footholds into new work on spider vision and the molecular basis of its extant variety. We therefore discuss what some of these first steps might be in the hopes of convincing others to join us in studying the vision of these fascinating creatures.

SUMMARYJumping spiders, or salticids, sample their environment using a combination of two types of eyes. The forward-facing pair of 'principal' eyes have narrow fields of view, but exceptional spatial resolution, while the two or three pairs of 'secondary' eyes have wide fields of view and function especially well as motion analysers. Motion detected by the secondary eyes may elicit an orienting response, whereupon the object of interest is examined further using the high-acuity principal eyes. The anterior lateral (AL) eyes are particularly interesting, as they are the only forward-facing pair of secondary eyes. In this study, we aimed to determine characteristics of stimuli that elicit orienting responses mediated by the AL eyes. After covering all eyes except the AL eyes, we measured orienting responses to dot stimuli that varied in size and contrast, and moved at different speeds. We found that all stimulus parameters had significant effects on orientation propensity. When tethered flies were used as prey, we found that visual information from the AL eyes alone was sufficient to elicit stalking behaviour. These results suggest that, in terms of overall visual processing, the relevance of spatial vision in the AL eyes has been underestimated in the literature. Our results also show that female spiders are significantly more responsive than males. We found that hunger caused similar increases in orientation propensity in the two sexes, but females responded more often than males both when sated and when hungry. A higher propensity by females to orient toward moving objects may be related to females tending to experience higher nutritional demands than males.

Jumping spiders (Salticidae) are renowned for their high performing visual system. In addition to their prominent forward-facing telescope-like principal eyes, salticids possess two or three pairs of secondary eyes used for wide-angle motion detection. Salticids orient towards relevant sources of motion detected by the secondary eyes, enabling them to inspect the stimulus with their spatially acute principal eyes. The anteriormost pair of secondary eyes, the anterior lateral (AL) eyes, also faces forward and has higher spatial acuity than the other, laterally-facing, secondary eyes. We used small computer-generated targets to elicit orienting saccades from tethered jumping spiders in order to examine the perceptual limits of the AL eyes. We describe the contrast thresholds of male and female spiders, investigate the reaction time between stimulus appearance and initiation of orientation, as well as the minimum distance a stimulus must travel before eliciting a saccade. Our results show that female spiders react to lower contrast stimuli than males and demonstrate that the secondary eyes can detect stimulus displacements considerably smaller than the inter-receptor angle.

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