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PubMed2005

In search of an auditory engram.

Monkeys trained preoperatively on a task designed to assess auditory recognition memory were impaired after removal of either the rostral superior temporal gyrus or the medial temporal lobe but were unaffected by lesions of the rhinal cortex. Behavioral analysis indicated that this result occurred because the monkeys did not or could not use long-term auditory recognition, and so depended instead on short-term working memory, which is unaffected by rhinal lesions. The findings suggest that monkeys may be unable to place representations of auditory stimuli into a long-term store and thus question whether the monkey's cerebral memory mechanisms in audition are intrinsically different from those in other sensory modalities. Furthermore, it raises the possibility that language is unique to humans not only because it depends on speech but also because it requires long-term auditory memory.

Animals↗
PubMed2006

Cortical signatures of noun and verb production.

Categories like "noun" and "verb" represent the basic units of grammar in all human languages, and the retrieval of categorical information associated with words is an essential step in the production of grammatical speech. Studies of brain-damaged patients suggest that knowledge of nouns and verbs can be spared or impaired selectively; however, the neuroanatomical correlates of this dissociation are not well understood. We used event-related functional MRI to identify cortical regions that were active when English-speaking subjects produced nouns or verbs in the context of short phrases. Two regions, in the left prefrontal cortex and left superior parietal lobule, were selectively activated for verb trials compared with noun trials; one region in the left inferior temporal lobe was more active during noun production than verb production. We propose that these regions are involved in representing core conceptual properties of nouns and verbs.

Adult↗
PubMed2006

Wnt3a gradient converts radial to bilateral feather symmetry via topological arrangement of epithelia.

The evolution of bilaterally symmetric feathers is a fundamental process leading toward flight. One major unsolved mystery is how the feathers of a single bird can form radially symmetric downy feathers and bilaterally symmetric flight feathers. In developing downy feather follicles, barb ridges are organized parallel to the long axis of the feather follicle. In developing flight-feather follicles, the barb ridges are organized helically toward the anterior region, leading to the fusion and creation of a rachis. Here we discover an anterior-posterior molecular gradient of wingless int (Wnt3)a in flight but not downy feathers. Global inhibition of the Wnt gradient transforms bilaterally symmetric feathers into radially symmetric feathers. Production of an ectopic local Wnt3a gradient reoriented barb ridges toward the source and created an ectopic rachis. We further show that the orientation of the Wnt3a gradient is dictated by the dermal papilla (DP). Swapping DPs between wing covert and breast downy feathers demonstrates that both feather symmetry and molecular gradients are in accord with the origin of the DP. Thus the fates of feather epidermal cells are not predetermined through some molecular codes but can be modulated. Together, our data suggest feathers are shaped by a DP--> Wnt gradient-->helical barb ridge organization-->creation of rachis-->bilateral symmetry sequence. We speculate diverse feather forms can be achieved by adjusting the orientation and slope of molecular gradients, which then shape the topological arrangements of feather epithelia, thus linking molecular activities to organ forms and novel functions.

Animals↗
PubMed2006

Organization of spines on the dendrites of Purkinje cells.

Dendritic spines have been investigated intensively over recent years; however, little is yet known about how they organize on the cell surface to make synaptic contacts with appropriate axons. Here we investigate spine distributions along the distal dendrites of cerebellar Purkinje cells, after biolistic labeling of intact tissue with a lipid-soluble dye. We show that the spines have a preference to form regular linear arrays and to trace short-pitch helical paths. The helical ordering is not determined by external factors that may influence how individual spines develop, because the same periodicities were present in fish and mammalian Purkinje cells, including those of weaver mice, which are depleted of the normal presynaptic partners for the spines. The ordering, therefore, is most likely an inherent property of the dendrite. Image reconstruction of dendrites from the different tissues showed that the helical spine distributions invariably lead to approximately equal sampling of surrounding space by the spineheads. The purpose of this organization may therefore be to maximize the opportunity of different spines to interact with different axons.

Animals↗
PubMed2005

Modeling the spatiotemporal cortical activity associated with the line-motion illusion in primary visual cortex.

Our large-scale computational model of the primary visual cortex that incorporates orientation-specific, long-range couplings with slow NMDA conductances operates in a fluctuating dynamic state of intermittent desuppression (IDS), which captures the behavior of coherent spontaneous cortical activity, as revealed by in vivo optical imaging based on voltage-sensitive dyes. Here, we address the functional significance of the IDS cortical operating points by investigating our model cortex response to the Hikosaka line-motion illusion (LMI) stimulus-a cue of a quickly flashed stationary square followed a few milliseconds later by a stationary bar. As revealed by voltage-sensitive dye imaging, there is an intriguing similarity between the cortical spatiotemporal activity in response to (i) the Hikosaka LMI stimulus and (ii) a small moving square. This similarity is believed to be associated with the preattentive illusory motion perception. Our numerical cortex produces similar spatiotemporal patterns in response to the two stimuli above, which are both in very good agreement with experimental results. The essential network mechanisms underpinning the LMI phenomenon in our model are (i) the spatiotemporal structure of the LMI input as sculpted by the lateral geniculate nucleus, (ii) a priming effect of the long-range NMDA-type cortical coupling, and (iii) the NMDA conductance-voltage correlation manifested in the IDS state. This mechanism in our model cortex, in turn, suggests a physiological underpinning for the LMI-associated patterns in the visual cortex of anaesthetized cat.

Animals↗
PubMed2006

A gain-control theory of binocular combination.

In binocular combination, light images on the two retinas are combined to form a single "cyclopean" perceptual image, in contrast to binocular rivalry which occurs when the two eyes have incompatible ("rivalrous") inputs and only one eye;s stimulus is perceived. We propose a computational theory for binocular combination with two basic principles of interaction: in every spatial neighborhood, each eye (i) exerts gain control on the other eye's signal in proportion to the contrast energy of its own input and (ii) additionally exerts gain control on the other eye's gain control. For stimuli of ordinary contrast, when either eye is stimulated alone, the predicted cyclopean image is the same as when both eyes are stimulated equally, coinciding with an easily observed property of natural vision. The gain-control theory is contrast dependent: Very low-contrast stimuli to the left- and right-eye add linearly to form the predicted cyclopean image. The intrinsic nonlinearity manifests itself only as contrast increases. To test the theory more precisely, a horizontal sine wave grating of 0.68 cycles per degree is presented to each eye. The gratings differ in contrast and phase. The predicted (and perceived) cyclopean grating also is a sine wave; its apparent phase indicates the relative contribution of the two eyes to the cyclopean image. For 48 measured combinations of phase and contrast, the theory with only one estimated parameter accounts for 95% of the variance of the data. Therefore, a simple, robust, physiologically plausible gain-control theory accurately describes an early stage of binocular combination.

Contrast Sensitivity↗
PubMed2006

Molecular evidence for deep evolutionary roots of bilaterality in animal development.

Nearly all metazoans show signs of bilaterality, yet it is believed the bilaterians arose from radially symmetric forms hundreds of millions of years ago. Cnidarians (corals, sea anemones, and "jellyfish") diverged from other animals before the radiation of the Bilateria. They are diploblastic and are often characterized as being radially symmetrical around their longitudinal (oral-aboral) axis. We have studied the deployment of orthologs of a number of family members of developmental regulatory genes that are expressed asymmetrically during bilaterian embryogenesis from the sea anemone, Nematostella vectensis. The secreted TGF-beta genes Nv-dpp, Nv-BMP5-8, six TGF-beta antagonists (NvChordin, NvNoggin1, NvNoggin2, NvGremlin, NvFollistatin, and NvFollistatin-like), the homeodomain proteins NvGoosecoid (NvGsc) and NvGbx, and the secreted guidance factor, NvNetrin, were studied. NvDpp, NvChordin, NvNoggin1, NvGsc, and NvNetrin are expressed asymmetrically along the axis perpendicular to the oral-aboral axis, the directive axis. Furthermore, NvGbx, and NvChordin are expressed in restricted domains on the left and right sides of the body, suggesting that the directive axis is homologous with the bilaterian dorsal-ventral axis. The asymmetric expression of NvNoggin1 and NvGsc appear to be maintained by the canonical Wnt signaling pathway. The asymmetric expression of NvNoggin1, NvNetrin, and Hox orthologs NvAnthox7, NvAnthox8, NvAnthox1a, and NvAnthox6, in conjunction with the observation that NvNoggin1 is able to induce a secondary axis in Xenopus embryos argues that N. vectensis could possess antecedents of the organization of the bilaterian central nervous system.

Animals↗
PubMed2006

Theoretical model of temperature regulation in the brain during changes in functional activity.

The balance between metabolic heat production, heat removal by blood flow, and heat conductance defines local temperature distribution in a living tissue. Disproportional local increases in blood flow as compared with oxygen consumption during functional brain activity disturb this balance, leading to temperature changes. In this article we have developed a theoretical framework that allows analysis of temperature changes during arbitrary functional brain activity. We established theoretical boundaries on temperature changes and explained how these boundaries depend on physiology (blood flow and metabolism) and external (heat exchange with the environment) experimental conditions. We show that, in regions located deep in the brain, task performance should be accompanied by temperature decreases in regions where blood flow increases (activated regions) and by temperature increases in regions where blood flow decreases (deactivated regions). The sign of temperature effect may be reversed for superficial cortex regions, where the baseline brain temperature is lower than the temperature of incoming arterial blood due to the heat exchange with the environment. Importantly, due to heat conductance, the temperature effect is not localized to the activated region but extends to a surrounding tissue at rest over the distances regulated by the temperature-shielding effect of blood flow. This temperature-shielding effect quantifies the means by which cerebral blood flow prevents "temperature perturbations" from propagating away from the perturbed regions. For small activated regions, this effect also substantially suppresses the magnitude of the temperature response, making it especially important for small animal brains.

Animals↗
PubMed2003

An egalitarian network model for the emergence of simple and complex cells in visual cortex.

We explain how simple and complex cells arise in a large-scale neuronal network model of the primary visual cortex of the macaque. Our model consists of approximately 4000 integrate-and-fire, conductance-based point neurons, representing the cells in a small, 1-mm(2) patch of an input layer of the primary visual cortex. In the model the local connections are isotropic and nonspecific, and convergent input from the lateral geniculate nucleus confers cortical cells with orientation and spatial phase preference. The balance between lateral connections and lateral geniculate nucleus drive determines whether individual neurons in this recurrent circuit are simple or complex. The model reproduces qualitatively the experimentally observed distributions of both extracellular and intracellular measures of simple and complex response.

Animals↗
PubMed1998

Statistical analysis of shape through triangulation of landmarks: A study of sexual dimorphism in hominids.

Two objects with homologous landmarks are said to be of the same shape if the configuration of landmarks of one object can be exactly matched with that of the other by translation, rotation/reflection, and scaling. In an earlier paper, the authors proposed statistical analysis of shape by considering logarithmic differences of all possible Euclidean distances between landmarks. Tests of significance for differences in the shape of objects and methods of discrimination between populations were developed with such data. In the present paper, the corresponding statistical methodology is developed by triangulation of the landmarks and by considering the angles as natural measurements of shape. This method is applied to the study of sexual dimorphism in hominids.

Animals↗
PubMed1999

Comparing in vitro, in situ, and in vivo experimental data in a three-dimensional model of mammalian cochlear mechanics.

Normal mammalian hearing is refined by amplification of the motion of the cochlear partition. This partition, comprising the organ of Corti sandwiched between the basilar and tectorial membranes, contains the outer hair cells that are thought to drive this amplification process. Force generation by outer hair cells has been studied extensively in vitro and in situ, but, to understand cochlear amplification fully, it is necessary to characterize the role played by each of the components of the cochlear partition in vivo. Observations of cochlear partition motion in vivo are severely restricted by its inaccessibility and sensitivity to surgical trauma, so, for the present study, a computer model has been used to simulate the operation of the cochlea under different experimental conditions. In this model, which uniquely retains much of the three-dimensional complexity of the real cochlea, the motions of the basilar and tectorial membranes are fundamentally different during in situ- and in vivo-like conditions. Furthermore, enhanced outer hair cell force generation in vitro leads paradoxically to a decrease in the gain of the cochlear amplifier during sound stimulation to the model in vivo. These results suggest that it is not possible to extrapolate directly from experimental observations made in vitro and in situ to the normal operation of the intact organ in vivo.

Acetylcholine↗
PubMed2005

Genetic ablation of polysialic acid causes severe neurodevelopmental defects rescued by deletion of the neural cell adhesion molecule.

Poly-alpha2,8-sialic acid (polySia) is a unique modification of the neural cell adhesion molecule, NCAM, tightly associated with neural development and plasticity. However, the vital role attributed to this carbohydrate polymer has been challenged by the mild phenotype of mice lacking polySia due to NCAM-deficiency. To dissect polySia and NCAM functions, we generated polySia-negative but NCAM-positive mice by simultaneous deletion of the two polysialyltransferase genes, St8sia-II and St8sia-IV. Beyond features shared with NCAM-null animals, a severe phenotype with specific brain wiring defects, progressive hydrocephalus, postnatal growth retardation, and precocious death was observed. These drastic defects were selectively rescued by additional deletion of NCAM, demonstrating that they originate from a gain of NCAM functions because of polySia deficiency. The data presented in this study reveal that the essential role of polySia resides in the control and coordination of NCAM interactions during mouse brain development. Moreover, this first demonstration in vivo that a highly specific glycan structure is more important than the glycoconjugate as a whole provides a novel view on the relevance of protein glycosylation for the complex process of building the vertebrate brain.

Animals↗
PubMed2006

Low conductance gap junctions mediate specific electrical coupling in body-wall muscle cells of Caenorhabditis elegans.

Invertebrate innexins and their mammalian homologues, the pannexins, are gap junction proteins. Although a large number of such proteins have been identified, few of the gap junctions that they form have been characterized to provide combined information of biophysical properties, coupling pattern, and molecular compositions. We adapted the dual whole cell voltage clamp technique to in situ analysis of electrical coupling in Caenorhabditis elegans body-wall muscle. We found that body-wall muscle cells were electrically coupled in a highly organized and specific pattern. The coupling was characterized by small (350 pS or less) junctional conductance (G(j)), which showed a bell-shaped relationship with junctional potential (V(j)) but was independent of membrane potential (V(m)). Injection of currents comparable to the junctional current (I(j)) into body-wall muscle cells caused significant depolarization, suggesting important functional relevance. The innexin UNC-9 appeared to be a key component of the gap junctions. Both Myc- and green fluorescent protein-tagged UNC-9 was localized to muscle intercellular junctions. G(j) was greatly inhibited in unc-9(fc16), a putative null mutant. Specific inhibition of UNC-9 function in muscle cells reduced locomotion velocity. Despite UNC-9 expression in both motor neurons and body-wall muscle cells, analyses of miniature and evoked postsynaptic currents in the unc-9 mutant showed normal neuromuscular transmission. These analyses provide a relatively detailed description of innexin-based gap junctions in a native tissue and suggest that innexin-based small conductance gap junctions can play an important role in processes such as locomotion.

Animals↗
PubMed2006

Evidence that the satin hair mutant gene Foxq1 is among multiple and functionally diverse regulatory targets for Hoxc13 during hair follicle differentiation.

It is increasingly evident that the molecular mechanisms underlying hair follicle differentiation and cycling recapitulate principles of embryonic patterning and organ regeneration. Here we used Hoxc13-overexpressing transgenic mice (also known as GC13 mice), known to develop severe hair growth defects and alopecia, as a tool for defining pathways of hair follicle differentiation. Gene array analysis performed with RNA from postnatal skin revealed differential expression of distinct subsets of genes specific for cells of the three major hair shaft compartments (cuticle, cortex, and medulla) and their precursors. This finding correlates well with the structural defects observed in each of these compartments and implicates Hoxc13 in diverse pathways of hair follicle differentiation. The group of medulla-specific genes was particularly intriguing because this included the developmentally regulated transcription factor-encoding gene Foxq1 that is altered in the medulladefective satin mouse hair mutant. We provide evidence that Foxq1 is a downstream target for Hoxc13 based on DNA binding studies as well as co-transfection and chromatin immunoprecipitation assays. Expression of additional medulla-specific genes down-regulated upon overexpression of Hoxc13 requires functional Foxq1 as their expression is ablated in hair follicles of satin mice. Combined, these results demonstrate that Hoxc13 and Foxq1 control medulla differentiation through a common regulatory pathway. The apparent regulatory interactions between members of the mammalian Hox and Fox gene families shown here may establish a paradigm for "cross-talk" between these two conserved regulatory gene families in different developmental contexts including embryonic patterning as well as organ development and renewal.

3T3 Cells↗
PubMed2001

Overview of mathematical computer models of striated sphincter muscles.

The present paper compares current mathematical striated sphincter models. Current models are subdivided in four categories: (1) simple models, (2) implementations of the urethral resistance relation, (3) models with realistic muscle dynamics and (4) finite element models. In our research group a neural network model, representing Onuf's nucleus, the spinal motor nucleus that innervates the external urethral and anal sphincters, was developed. A realistic sphincter model is needed to test the neural network. To decide whether or not a model is applicable in our research two requirements should be fulfilled: (1) the presence of realistic muscle dynamics preferably by implementation of a Huxley type muscle model and (2) the model should consist of more than one muscle unit to form a more dimensional model. Reviewing the literature, if a myogenic sphincter is modelled, mainly the Hill-equation is applied. Moreover, single muscle unit models are published. In general a multi-unit muscle model of the sphincter is lacking, prohibiting the study of the inherent properties of sphincter muscles, which could give information on the realistic behaviour of elements in circular muscles. It is concluded that the functionality of current sphincter models is limited for our purpose.

Animals↗
PubMed2003

Revised formulas for summarizing retinal vessel diameters.

Background/Purpose. Recent findings suggest that an objective assessment of retinal vessel caliber from fundus photographs provide information about the association of microvascular characteristics with macrovascular disease. Current methods used to quantify retinal vessel caliber, introduced by Parr(1,2) and Hubbard,(3) are not independent of scale and are affected by the number of vessels. To improve upon these methods we introduce revised formulas for quantifying vessel caliber. Methods. Revised formulas were estimated using retinal vessel measurements from 44 young adults free of hypertension and diabetes. Comparisons between the two methods were done using digitized photographs from 4926 participants at the baseline examination of the Beaver Dam Eye Study (BDES), an ongoing population-based cohort study initiated in 1987. Individual arterioles and venules were measured using semi-automated computer software from which summary measures were calculated. Results. Correlation coefficients between the Parr-Hubbard and revised formulas were high (Pearson correlation coefficients ranging from 0.94 to 0.98). Both arteriolar and venular caliber significantly increased with an increasing number of vessels measured using the Parr-Hubbard formulas (p < 0.001), which in turn affected the relationship to mean arterial blood pressure. To the contrary, the revised formulas were not affected by the number of measured vessels (p > 0.50). Conclusions. We describe revised formulas for summarizing retinal vessel diameters measured from fundus photographs to be used in future studies and analyses. The revised formulas correlate highly with the previously used Parr-Hubbard formulas, but offer the advantages of being more robust against variability in the number of vessels observed, being independent of image scale, and being easier to implement.

Adult↗
PubMed2003

Higher-order wavefront aberrations induced by small ablation area and sub-clinical decentration in simulated corneal refractive surgery using a perturbed schematic eye model.

The purpose was to investigate the effect of decentered ablation in myopic corneal refractive surgery by simulation. Wavefront aberrations in a small pupil area (radius: 2 mm) for photopic vision and in a large pupil area (radius: 3 mm) for mesopic vision with simulated refractive surgery were analyzed using Zernike polynomials. Radii of ablation were 3, 2.5 and 2 mm. Decentrations of ablation were 0 mm and 0.5 mm. Change of the surface shape by ablation was considered a perturbation, while Gullstrand's schematic eye was used as un-perturbed optics. For photopic vision, wavefront aberrations were about the same as with un-perturbed optics. For mesopic vision, the results were heavily dependent on the radius of ablation area. When the radius was 3 mm, wavefront aberrations did not increase very much compared to un-perturbed optics. When the radius was smaller than 3 mm, spherical aberration was induced by centered ablations, and coma was also induced by decentered ablations. In conclusion, small ablation areas or subclinical decentrations of ablations could cause serious amounts of wavefront aberrations to the optics of the eye in the simulations.

Cornea↗
PubMed2003

Dynamic eye plant models and the control of eye movements.

Models of the oculomotor plant (globe, muscles, pulleys, and orbital tissues) fall into three categories: 1). one-dimensional dynamic with lumped plant elements, 2). three-dimensional dynamic with lumped plant elements, or 3). three-dimensional static with distinct plant elements. The second class of models is most often used when studying the neural control of 3-D eye movement, because they best represent the plant dynamics. However, they are often faulted because they make two unrealistic assumptions: 1). muscle pairs act along the three orthogonal axes (symmetry assumption); and 2). the force generated by the muscles depends only on their innervation (force assumption). It turns out that the symmetry assumption is quite benign, because in a realistic model of the plant the deviations from orthogonal axes can be easily accounted for by simple adjustments to the innervation. In contrast, the force assumption introduces some serious problems. In the present paper, the authors show that a realistic, dynamic model of the geometry of the orbit, with independent muscles, makes different predictions than a similar model with lumped muscles. This difference arises because muscle force is a function of both innervation and muscle length.

Biomechanical Phenomena↗