Does experience provide a permissive or instructive influence on the development of direction selectivity in visual cortex?

In principle, the development of sensory receptive fields in cortex could arise from experience-independent mechanisms that have been acquired through evolution, or through an online analysis of the sensory experience of the individual animal. Here we review recent experiments that suggest that the development of direction selectivity in carnivore visual cortex requires experience, but also suggest that the experience of an individual animal cannot greatly influence the parameters of the direction tuning that emerges, including direction angle preference and speed tuning. The direction angle preference that a neuron will acquire can be predicted from small initial biases that are present in the naïve cortex prior to the onset of visual experience. Further, experience with stimuli that move at slow or fast speeds does not alter the speed tuning properties of direction-selective neurons, suggesting that speed tuning preferences are built in. Finally, unpatterned optogenetic activation of the cortex over a period of a few hours is sufficient to produce the rapid emergence of direction selectivity in the naïve ferret cortex, suggesting that information about the direction angle preference that cells will acquire must already be present in the cortical circuit prior to experience. These results are consistent with the idea that experience has a permissive influence on the development of direction selectivity.

Erratum to: S-wave attenuation in northeastern Sonora, Mexico, near the faults that ruptured during the earthquake of 3 May 1887 Mw 7.5.

[This corrects the article DOI: 10.1186/2193-1801-3-747.].

Emerging feed-forward inhibition allows the robust formation of direction selectivity in the developing ferret visual cortex.

The computation of direction selectivity requires that a cell respond to joint spatial and temporal characteristics of the stimulus that cannot be separated into independent components. Direction selectivity in ferret visual cortex is not present at the time of eye opening but instead develops in the days and weeks following eye opening in a process that requires visual experience with moving stimuli. Classic Hebbian or spike timing-dependent modification of excitatory feed-forward synaptic inputs is unable to produce direction-selective cells from unselective or weakly directionally biased initial conditions because inputs eventually grow so strong that they can independently drive cortical neurons, violating the joint spatial-temporal activation requirement. Furthermore, without some form of synaptic competition, cells cannot develop direction selectivity in response to training with bidirectional stimulation, as cells in ferret visual cortex do. We show that imposing a maximum lateral geniculate nucleus (LGN)-to-cortex synaptic weight allows neurons to develop direction-selective responses that maintain the requirement for joint spatial and temporal activation. We demonstrate that a novel form of inhibitory plasticity, postsynaptic activity-dependent long-term potentiation of inhibition (POSD-LTPi), which operates in the developing cortex at the time of eye opening, can provide synaptic competition and enables robust development of direction-selective receptive fields with unidirectional or bidirectional stimulation. We propose a general model of the development of spatiotemporal receptive fields that consists of two phases: an experience-independent establishment of initial biases, followed by an experience-dependent amplification or modification of these biases via correlation-based plasticity of excitatory inputs that compete against gradually increasing feed-forward inhibition.

S-wave attenuation in northeastern Sonora, Mexico, near the faults that ruptured during the earthquake of 3 May 1887 Mw 7.5.

We used a new data set of relocated earthquakes recorded by the Seismic Network of Northeastern Sonora, Mexico (RESNES) to characterize the attenuation of S-waves in the fault zone of the 1887 Sonora earthquake (M w 7.5). We determined spectral attenuation functions for hypocentral distances (r) between 10 and 140 km using a nonparametric approach and found that in this fault zone the spectral amplitudes decay slower with distance at low frequencies (f < 4 Hz) compared to those reported in previous studies in the region using more distant recordings. The attenuation functions obtained for 23 frequencies (0.4 ≤ f ≤ 63.1 Hz) permit us estimating the average quality factor Q S = (141 ± 1.1 )f ((0.74 ± 0.04)) and a geometrical spreading term G(r) = 1/r (0.21). The values of Q estimated for S-wave paths traveling along the fault system that rupture during the 1887 event, in the north-south direction, are considerably lower than the average Q estimated using source-station paths from multiple stations and directions. These results indicate that near the fault zone S waves attenuate considerably more than at regional scale, particularly at low frequencies. This may be the result of strong scattering near the faults due to the fractured upper crust and higher intrinsic attenuation due to stress concentration near the faults.

Statistical traces of long-term memories stored in strengths and patterns of synaptic connections.

Learning and long-term memory rely on plasticity of neural circuits. In adult cerebral cortex, plasticity can result from potentiation and depression of synaptic strengths and structural reorganization of circuits through growth and retraction of dendritic spines. By analyzing 166 distributions of spine head volumes and spine lengths from mouse, rat, monkey, and human brains, we determine the "generalized cost" of dendritic spines. This cost universally depends on spine shape, i.e., the dependence is the same in all the analyzed systems. We show that, in adult, synaptic strength and structural synaptic plasticity mechanisms are in statistical equilibrium, the numbers of dendritic spines in different cortical areas are nearly optimally chosen for memory storage, and the distributions of spine lengths and head volumes are governed by a single parameter--the effective temperature. We suggest that the effective temperature may be viewed as a measure of circuit stability or longevity of stored memories.

Structural plasticity of circuits in cortical neuropil.

Learning and memory formation in the brain depend on the plasticity of neural circuits. In the adult and developing cerebral cortex, this plasticity can result from the formation and elimination of dendritic spines. New synaptic contacts appear in the neuropil where the gaps between axonal and dendritic branches can be bridged by dendritic spines. Such sites are termed potential synapses. Here, we describe a theoretical framework for the analysis of spine remodeling plasticity. We provide a quantitative description of two models of spine remodeling in which the presence of a bouton is either required or not for the formation of a new synapse. We derive expressions for the density of potential synapses in the neuropil, the connectivity fraction, which is the ratio of actual to potential synapses, and the number of structurally different circuits attainable with spine remodeling. We calculate these parameters in mouse occipital cortex, rat CA1, monkey V1, and human temporal cortex. We find that, on average, a dendritic spine can choose among 4-7 potential targets in rodents and 10-20 potential targets in primates. The potential of neuropil for structural circuit remodeling is highest in rat CA1 (7.1-8.6 bits/mum(3)) and lowest in monkey V1 (1.3-1.5 bits/mum(3)). We also evaluate the lower bound of neuron selectivity in the choice of synaptic partners. Postsynaptic excitatory neurons in rodents make synaptic contacts with >21-30% of presynaptic axons encountered with new spine growth. Primate neurons appear to be more selective, making synaptic connections with >7-15% of encountered axons.