Cephalopod Eyes excerpt from http://archives.evergreen.edu/webpages/curricular/20112012/m2o1112/web/cephalopods.html Camera-type Eye The cephalopods possess camera-type eyes similar to those of vertebrates. These eyes contain an iris, a rectangular pupil, a nearly circular lens, vitreous cavity and photoreceptors forming a retina. The pupil shape for different cephalopods varies. The octopus has a rectangular pupil, the cuttlefish a 'W' shaped pupil and the squid a circular pupil. They use their pupils to control the amount of light that hits the retina. The cephalopod eye resembles the vertebrate eye but is less sophisticated. Of all the cephalopods, the octopus has the most developed eye with a completely closed cornea. The octopus focuses by moving its entire (stiff) lens back and forth within the eye. Human use small muscles to change the shape of the lens and its distance to the retina. Considering that cephalopods are water-dwelling creatures, one of their most interesting features is their ability to keep their eye orientation the same in relation to gravity regardless of their body position in the three dimensional water column. Essentially, their pupil will stay horizontally aligned regardless of the angle of their body. In order for this to work, they have a balance organ called a statocyst that controls the optic muscles. A statocyst is a sense organ that is half filled with both tiny calcium granules in a jelly-like fluid and lined with hair cells. The statocyst gives the cephalopod input about orientation and acceleration similar to the utricle and saccule in human ears. Evolutionary Significance The likeness between the camera-type cephalopod eye and the vertebrate eye has been used as a classic example of convergent evolution for almost 140 years. Convergent evolution is the independent evolution of similar structural or functional components in two or more unrelated or distantly related lineages. The other possible explanation is parallel evolution wherein two lineages evolve from a common ancestor and in a common trajectory towards a similar endpoint, resulting in the similar or shared trait in different species. The convergence theory of the camera-type cephalopod eye is now being challenged. A conserved gene network has been discovered which includes the gene Pax6. Pax6 governs eye development in both vertebrate and invertebrate lineages. This gene conservation may indicate a common origin of eyes in vertebrates and invertebrates. If this is the case then the similarity between the camera-type cephalopod eye and the vertebrate eye would be a result of constraint in development and parallel evolution. However, it is both possible and probable that this likeness is the result of a combination of both convergent and parallel evolution, as the eye is a complex organ with many components which may evolve independently of one another.