25The retinal output is the sole source of visual information for the brain. Studies in non-primate 26 mammals estimate that this information is carried by several dozens of retinal ganglion cell 27 types, each informing the brain about different aspects of a visual scene. Even though 28 morphological studies of primate retina suggest a similar diversity of ganglion cell types, 29 research has focused on the function of only a few cell types. In human retina, recordings 30 from individual cells are anecdotal. Here, we present the first systematic ex-vivo recording of 31 light responses from 342 ganglion cells in human retinas obtained from donors. We find a 32 great variety in the human retinal output in terms of preferences for positive or negative 33 contrast, spatio-temporal frequency encoding, contrast sensitivity, and speed tuning. Some 34 human ganglion cells showed similar response behavior as known cell types in other primates, 35 while we also recorded light responses that have not been described previously. This first 36 extensive description of the human retinal output should facilitate interpretation of primate 37 data and comparison to other mammalian species, and it lays the basis for the use of ex-vivo 38 human retina for in-vitro analysis of novel treatment approaches.
40Vision starts in the retina, a highly structured part of the central nervous system. The retina 41 performs important signal processing: the incoming images are captured by the 42 photoreceptors, analyzed and split into parallel information streams by retinal circuits, and 43 sent along the optic nerve to higher visual brain centers. Each of the parallel information 44 streams is embodied by a type of ganglion cell and informs the brain about a particular aspect 45 of the visual scene 1 . The non-primate mammalian retina contains over 40 of these different 46 information streams, which can be distinguished based on both functional and morphological 47 criteria 2-7 .
48One striking feature of retinal architecture is that each ganglion cell type tiles the retina so 49 that each feature can be extracted at each location in the visual field. Nevertheless, regional 50 specializations do exist, for example the fovea of the primate retina, a region of very high 51 visual acuity. The foveal region consists almost exclusively of four retinal ganglion cell types, 52 the ON and OFF parasol cells and the ON and OFF midget cells 8-10 , which account for 50-53 70% of all ganglion cells in the primate retina 11 . Functional studies using non-human 54 primates have often focused on these four most abundant retinal ganglion cell types 12-17 . 55 Morphological studies of the complete primate retina, on the other hand, describe a similar 56 variety in ganglion cell types as found in the non-primate retina with at least 17 57 morphologically identified types 11,18-21 . However, functional studies of these non-foveal 58 ganglion cell types in non-human primates have been limited to a set of 7 types 14,22-27 , and 59 physiological assessment of t...