Optical Imaging of Retinotopic Maps in a Small Songbird,the Zebra Finch

Background

The primary visual cortex of mammals is characterised by a retinotopic representation of the visual field. It has therefore been speculated that the visual wulst, the avian homologue of the visual cortex, also contains such a retinotopic map. We examined this for the first time by optical imaging of intrinsic signals in zebra finches, a small songbird with laterally placed eyes. In addition to the visual wulst, we visualised the retinotopic map of the optic tectum which is homologue to the superior colliculus in mammals.

Methodology/Principal Findings

For the optic tectum, our results confirmed previous accounts of topography based on anatomical studies and conventional electrophysiology. Within the visual wulst, the retinotopy revealed by our experiments has not been illustrated convincingly before. The frontal part of the visual field (0°±30° azimuth) was not represented in the retinotopic map. The visual field from 30°–60° azimuth showed stronger magnification compared with more lateral regions. Only stimuli within elevations between about 20° and 40° above the horizon elicited neuronal activation. Activation from other elevations was masked by activation of the preferred region. Most interestingly, we observed more than one retinotopic representation of visual space within the visual wulst, which indicates that the avian wulst, like the visual cortex in mammals, may show some compartmentation parallel to the surface in addition to its layered structure.

Conclusion/Significance

Our results show the applicability of the optical imaging method also for small songbirds. We obtained a more detailed picture of retinotopic maps in birds, especially on the functional neuronal organisation of the visual wulst. Our findings support the notion of homology of visual wulst and visual cortex by showing that there is a functional correspondence between the two areas but also raise questions based on considerable differences between avian and mammalian retinotopic representations.     

Topographical Estimation of Visual Population Receptive Fields by fMRI

Visual cortex is retinotopically organized so that neighboring populations of cells map to neighboring parts of the visual field. Functional magnetic resonance imaging allows us to estimate voxel-based population receptive fields (pRF), i.e., the part of the visual field that activates the cells within each voxel. Prior, direct, pRF estimation methods¹ suffer from certain limitations: 1) the pRF model is chosen a-priori and may not fully capture the actual pRF shape, and 2) pRF centers are prone to mislocalization near the border of the stimulus space. Here a new topographical pRF estimation method² is proposed that largely circumvents these limitations. A linear model is used to predict the Blood Oxygen Level-Dependent (BOLD) signal by convolving the linear response of the pRF to the visual stimulus with the canonical hemodynamic response function. PRF topography is represented as a weight vector whose components represent the strength of the aggregate response of voxel neurons to stimuli presented at different visual field locations. The resulting linear equations can be solved for the pRF weight vector using ridge regression³, yielding the pRF topography. A pRF model that is matched to the estimated topography can then be chosen post-hoc, thereby improving the estimates of pRF parameters such as pRF-center location, pRF orientation, size, etc. Having the pRF topography available also allows the visual verification of pRF parameter estimates allowing the extraction of various pRF properties without having to make a-priori assumptions about the pRF structure. This approach promises to be particularly useful for investigating the pRF organization of patients with disorders of the visual system.     

Cambrian view

The analysis of visual systems is a valuable method of assessing phylogenetic processes. As in the present animal world, we find simple and complex systems in the Lower Cambrian. One may detect “simple eyes” for example with an advanced design in lobopodians, while the existence of even more simple “simple eyes” is very probable but still to be proved. More complex systems are to be found. In Leanchoilia illecebrosa Hou, 1987 and Leanchoilia superlata Walcott, 1912 there are probable dorsal median eyes and a pair of fine, stalked ventral eyes. Both systems may contribute to phylogenetic and systematic discussions. These presumably movable stalked eyes may be regarded as an adaptation to a mobile lifestyle. They suggest that the physiologic principle of nystagmus to stabilise the visual world of an animal in motion was already realised in Leanchoilia, perhaps for the first time. To analyse the surface of the early eyes from the Lower Cambrian – not only of Leanchoilia, but of any other forms as well – the number, shape and other parameters of the lenses could lead to further knowledge regarding vision in early invertebrates.     

Helical Perturbations of the Flagellar Filament:Rhizobium lupiniH13–3 at 13 Å Resolution

Flagellar filaments are highly conserved structures in terms of the underlying symmetry of the polymer, subunit domain organization of the flagellin monomer, amino acid composition and primary sequence at the N and C termini. Traditionally, filaments are classified as “plain” or “complex.” In complex filaments, the helical lattice is perturbed in a pairwise manner such that the symmetry is reduced along the 6-start helical lines. Both plain (unperturbed) and complex (helically perturbed) components are helically symmetric and share a common lattice. The perturbation inRhizobium lupiniH13–3 results in a subunit composed of a dimer of flagellin. We have generated a ≈13 Å resolution three-dimensional density map of the complex filament ofR. lupiniH13–3 from low-dose images of negatively stained filaments. Compared to a previous map, which extended to only ≈25 Å resolution and which was generated from only five filaments containing six layer-lines each, the current map is a product of merging 139 data sets containing 66 layer-lines each. The higher resolution and improved signal-to-noise yield a detailed and interpretable density map. The density map is divided into four concentric rings. These amount to two dense cylinders interconnected by low density radial spokes and wrapped by a three-start external winding. The unperturbed component of the map is strikingly similar to the known plain filament maps and, in particular, to that ofCaulobacter crescentus. The helically perturbed component contributes mainly to the filaments's exterior (domain D3) where it comprises the tips of the outer domains interconnecting, pairwise, along the 11-start protofilaments and, again, laterally along the 6-start lines forming vertical and horizontal loops. Strong intersubunit connectivity occurs in the D2 shell and in the inner shell which is dominated by 3-start densities. The contribution of the complex component to the radial spokes seems negligible.     

Evolution of eye development in arthropods: phylogenetic aspects

The architecture of the adult arthropod visual system for many decades has contributed important character sets that are useful for reconstructing the phylogenetic relationships within this group. In the current paper we explore whether aspects of eye development can also contribute new arguments to the discussion of arthropod phylogeny. We review the current knowledge on eye formation in Trilobita, Xiphosura, Myriapoda, Hexapoda, and Crustacea. All euarthropod taxa share the motif of a proliferation zone at the side of the developing eye field that contributes new eye elements. Two major variations of this common motif can be distinguished: 1. The “row by row type” of Trilobita, Xiphosura, and Diplopoda. In this type, the proliferation zone at the side of the eye field generates new single, large elements with a high and variable cell number, which are added to the side of the eye and extend rows of existing eye elements. Cell proliferation, differentiation and ommatidial assembly seem to be separated in time but spatially confined within the precursors of the optic units which grow continuously once they are formed (intercalary growth). 2. The “morphogenetic front type” of eye formation in Crustacea + Hexapoda (Tetraconata). In this type, there is a clear temporal and spatial separation of the formation and differentiation processes. Proliferation and the initial steps of pattern formation take place in linear and parallel mitotic and morphogenetic fronts (the mitotic waves and the morphogenetic furrow/transition zone) and numerous but small new elements with a strictly fixed set of cells are added to the eye field. In Tetraconata, once formed, the individual ommatidia do not grow any more. Scutigeromorph chilopods take an intermediate position between these two major types. We suggest that the “row by row type” as seen in Trilobita, Xiphosura and Diplopoda represents the plesiomorphic developmental mode of eye formation from the euarthropod ground pattern whereas the “morphogenetic front type” is apomorphic for the Tetraconata. Our data are discussed with regard to two competing hypotheses on arthropod phylogeny, the “Tracheata” versus “Tetraconata” concept. The modes of eye development in Myriapoda is more parsimonious to explain in the Tetraconata hypothesis so that our data raise the possibility that myriapod eyes may not be secondarily reconstructed insect eyes as the prevailing hypothesis suggests.     

Some nonstandard modeling techniques in theoretical biology

Three topics are surveyed. (1) The quasisteady-state assumption is a fundamental example of frequent instances in theoretical biology where advantage can be taken of the simultaneous existence of widely separated time scales. It is shown that appropriate scaling can provide a markedly sharper estimate of the range of validity of this approximation. (2) A new methodology for modelling dynamically changing networks is applied to simple morphogenesis in fungal colonies. A brief discussion is given of possible application to the complex polymerization and cross-linking that characterize cytoplasmic fibers. (3) It is shown that problems in ecology and immunology can be illuminated by considering nonlocal interactions (and hence integrodifferential equations) that generate patterns in “aspect space” or “shape space.”     

Forest reconstruction as ecological engineering

Land restoration involves reconstruction of the native biota in a sustainable form. If reconstruction involves deliberate manipulation of biological organisms and the physical-chemical environment to achieve specific human goals, it qualifies as ecological engineering. Restoration which uses natural processes to achieve endpoints which are unpredictable but can be accepted because they are “natural” is not ecological engineering. In Japan a system of forest reconstruction has been developed which is based on knowledge of the potential vegetation of a site, knowledge of the methods of germination and growth of the species which compose the mature vegetation and a method of site preparation and planting. This ecological engineering approach has been used on 285 sites, in a variety of habitats, to form dense stands of vegetation to hide industrial complexes, control visual, noise and chemical pollution, stabilize soil and beaches and provide urban green space. The technique has also been used to restore tropical rain forest.     

The phylogeny of the genus Ischioscia Verhoeff, 1928, with redescriptions of three species (Crustacea: Isopoda: Oniscidea)

The genus Ischioscia Verhoeff, 1928 is reviewed. 26 species are considered valid. A key for their identification is given, as well as a map showing the geographic distribution. The known range of the genus covers a large area from the central Amazon region to the mountains of Guatemala. The species of Ischioscia have a typical “philosciid” habitus (“runner” type); they can be distinguished from other Neotropical species with similar habitus by the following apomorphies: (1) male pereiopod 1 carpus enlarged to a plate-like extension, (2) scale field on male pereiopod 1 covering entire frontal side of the carpus, (3) male pereiopod 7 ischium with a ventral scale field, (4) dactylus in both sexes with a long inner claw. The groundpattern of Ischioscia is reconstructed, and an analysis of the phylogenetic relations within the genus is made on the basis of morphological data. The species are very similar to each other, most differences are found in the male structures of sexually dimorphic features. Ischioscia sturmi (Vandel, 1972), I. amazonica Lemos de Castro, 1955 and I. bolivari Vandel, 1968 are redescribed in detail.     

Retinal optic flow during natural locomotion

We examine the structure of the visual motion projected on the retina during natural locomotion in real world environments. Bipedal gait generates a complex, rhythmic pattern of head translation and rotation in space, so without gaze stabilization mechanisms such as the vestibular-ocular-reflex (VOR) a walker’s visually specified heading would vary dramatically throughout the gait cycle. The act of fixation on stable points in the environment nulls image motion at the fovea, resulting in stable patterns of outflow on the retinae centered on the point of fixation. These outflowing patterns retain a higher order structure that is informative about the stabilized trajectory of the eye through space. We measure this structure by applying the curl and divergence operations on the retinal flow velocity vector fields and found features that may be valuable for the control of locomotion. In particular, the sign and magnitude of foveal curl in retinal flow specifies the body’s trajectory relative to the gaze point, while the point of maximum divergence in the retinal flow field specifies the walker’s instantaneous overground velocity/momentum vector in retinotopic coordinates. Assuming that walkers can determine the body position relative to gaze direction, these time-varying retinotopic cues for the body’s momentum could provide a visual control signal for locomotion over complex terrain. In contrast, the temporal variation of the eye-movement-free, head-centered flow fields is large enough to be problematic for use in steering towards a goal. Consideration of optic flow in the context of real-world locomotion therefore suggests a re-evaluation of the role of optic flow in the control of action during natural behavior.     

Tracking down the "head blob": comparative analysis of wingless expression in the developing insect procephalon reveals progressive reduction of embryonic visual system patterning in higher insects

The evolution of larval head morphology in holometabolous insects is characterized by reduction of antennal appendages and the visual system components. Little insight has been gained into molecular developmental changes underlying this morphological diversification. Here we compare the expression of the segment polarity gene wingless (wg) in the pregnathal head of fruit fly, flour beetle and grasshopper embryos. We provide evidence that wg activity contributes to segment border formation, and, subsequently, the separation of the visual system and protocerebrum anlagen in the anterior procephalon. In directly developing insects like grasshopper, seven expression domains are formed during this process. The activation of four of these, which correspond to polar expression pairs in the optic lobe anlagen and the protocerebral ectoderm, has shifted to postembryonic stages in flour beetle and Drosophila. The remaining three domains map to the protocerebral neuroectoderm, and form by disintegration of a large precursor domain in flour beetle and grasshopper. In Drosophila, the precursor domain remains intact, constituting the previously described “head blob”. These data document major changes in the expression of an early patterning gene correlated with the dramatic evolution of embryonic visual system development in the Holometabola.     

Neural plasticity: how the eye tells the brain about sound location

Recent studies in owls and ferrets seem to have identified the origin and nature of the visual signals that shape the development of the auditory space map in the midbrain, which ensures that the neural representations of both sensory modalities share the same topographic organization.     

The coordinated mapping of visual space and response features in visual cortex

Whether general principles can explain the layouts of cortical maps remains unresolved. In primary visual cortex of ferret, the relationships between the maps of visual space and response features are predicted by a "dimension-reduction" model. The representation of visual space is anisotropic, with the elevation and azimuth axes having different magnification. This anisotropy is reflected in the orientation, ocular dominance, and spatial frequency domains, which are elongated such that their directions of rapid change, or high-gradient axes, are orthogonal to the high-gradient axis of the visual map. The feature maps are also strongly interdependent-their high-gradient regions avoid one another and intersect orthogonally where essential, so that overlap is minimized. Our results demonstrate a clear influence of the visual map on each feature map. In turn, the local representation of visual space is smooth, as predicted when many features are mapped within a cortical area.     

Construction of a DNA nano-object directly demonstrates computation

We demonstrate a computing method in which a DNA nano-object representing the solution of a problem emerges as a result of self-assembly. We report an experiment in which three-vertex colorability for a six-vertex graph with nine edges is solved by constructing a DNA molecule representing the colored graph itself. Our findings show that computation based on “shape processing” is a viable alternative to symbol processing when computing by molecular self-assembly.     

Features of the Retinotopic Representation in the Visual Wulst of a Laterally Eyed Bird,the Zebra Finch (Taeniopygia guttata)

The visual wulst of the zebra finch comprises at least two retinotopic maps of the contralateral eye. As yet, it is not known how much of the visual field is represented in the wulst neuronal maps, how the organization of the maps is related to the retinal architecture, and how information from the ipsilateral eye is involved in the activation of the wulst. Here, we have used autofluorescent flavoprotein imaging and classical anatomical methods to investigate such characteristics of the most posterior map of the multiple retinotopic representations. We found that the visual wulst can be activated by visual stimuli from a large part of the visual field of the contralateral eye. Horizontally, the visual field representation extended from -5° beyond the beak tip up to +125° laterally. Vertically, a small strip from -10° below to about +25° above the horizon activated the visual wulst. Although retinal ganglion cells had a much higher density around the fovea and along a strip extending from the fovea towards the beak tip, these areas were not overrepresented in the wulst map. The wulst area activated from the foveal region of the ipsilateral eye, overlapped substantially with the middle of the three contralaterally activated regions in the visual wulst, and partially with the other two. Visual wulst activity evoked by stimulation of the frontal visual field was stronger with contralateral than with binocular stimulation. This confirms earlier electrophysiological studies indicating an inhibitory influence of the activation of the ipsilateral eye on wulst activity elicited by stimulating the contralateral eye. The lack of a foveal overrepresentation suggests that identification of objects may not be the primary task of the zebra finch visual wulst. Instead, this brain area may be involved in the processing of visual information necessary for spatial orientation.     

Some remarks on the interpretation of Penrose's size and shape components. Craniometrical similarities of African groups

The present paper deals with cranial data of different African races and shows how far the components “size” and “shape” of the Penrose distance express biological distances as well as the kind of changes occurring when different groups of variables are used. The stability of the distances between the populations was further studied by exchanging the reference group on a racial and subracial level. The analyses supply a differentiated picture according to the affinities, especially between the reference populations of the major races. Europoid and Negroid populations can often be separated only by shape distance, while the size distance is more suitable for separating Bushmen and Hottentots.     

Complex Generalized Synchronization and Parameter Identification of Nonidentical Nonlinear Complex Systems

In this paper, generalized synchronization (GS) is extended from real space to complex space, resulting in a new synchronization scheme, complex generalized synchronization (CGS). Based on Lyapunov stability theory, an adaptive controller and parameter update laws are designed to realize CGS and parameter identification of two nonidentical chaotic (hyperchaotic) complex systems with respect to a given complex map vector. This scheme is applied to synchronize a memristor-based hyperchaotic complex Lü system and a memristor-based chaotic complex Lorenz system, a chaotic complex Chen system and a memristor-based chaotic complex Lorenz system, as well as a memristor-based hyperchaotic complex Lü system and a chaotic complex Lü system with fully unknown parameters. The corresponding numerical simulations illustrate the feasibility and effectiveness of the proposed scheme.