The Role of Ongoing Dendritic Oscillations in Single-Neuron Dynamics

The dendritic tree contributes significantly to the elementary computations a neuron performs while converting its synaptic inputs into action potential output. Traditionally, these computations have been characterized as both temporally and spatially localized. Under this localist account, neurons compute near-instantaneous mappings from their current input to their current output, brought about by somatic summation of dendritic contributions that are generated in functionally segregated compartments. However, recent evidence about the presence of oscillations in dendrites suggests a qualitatively different mode of operation: the instantaneous phase of such oscillations can depend on a long history of inputs, and under appropriate conditions, even dendritic oscillators that are remote may interact through synchronization. Here, we develop a mathematical framework to analyze the interactions of local dendritic oscillations and the way these interactions influence single cell computations. Combining weakly coupled oscillator methods with cable theoretic arguments, we derive phase-locking states for multiple oscillating dendritic compartments. We characterize how the phase-locking properties depend on key parameters of the oscillating dendrite: the electrotonic properties of the (active) dendritic segment, and the intrinsic properties of the dendritic oscillators. As a direct consequence, we show how input to the dendrites can modulate phase-locking behavior and hence global dendritic coherence. In turn, dendritic coherence is able to gate the integration and propagation of synaptic signals to the soma, ultimately leading to an effective control of somatic spike generation. Our results suggest that dendritic oscillations enable the dendritic tree to operate on more global temporal and spatial scales than previously thought; notably that local dendritic activity may be a mechanism for generating on-going whole-cell voltage oscillations.     

Changes in sperm stores, ejaculate size, fertilization success, and sexual motivation over repeated matings in the common toad, Bufo bufo (Anura: Bufonidae)

Fertilization is of central importance in the determination of reproductive success for both males and females. In species where males have the chance to mate repeatedly within a short period of time, sperm stocks may become depleted and males may have to carefully economize on their sperm reserves. Also, intensive intrasexual competition for females and repeated matings may lead to exhaustion on the behavioural level. To determine whether the reproductive potential of males is limited and if such a limitation is due to behavioural exhaustion or sperm depletion, we experimentally investigated changes in sperm stores, sperm expenditure, fertilization success, and sexual motivation over three repeated matings in the common toad, Bufo bufo , where the breeding season is short and sequential polygyny occurs. At the end of the breeding season, the number of sperm stored in the testes of males mated repeatedly was close to 50% lower than in testes of unmated males. Ejaculate size, which was estimated by applying a novel method allowing direct quantification, decreased by 88% from first to third matings. We also observed a drop in fertilization success from the first two to third matings by 65%, which was largest in males that had started the reproductive season in bad body condition. Some of these males also showed a decreased interest in females in the third mating round. Our results suggest that sperm depletion and loss of sexual motivation may together set a limit to the reproductive potential of common toad males. The present study draws attention to a limitation in reproductive potential, which may occur more often than currently anticipated and has the potential to strongly influence several aspects of reproductive behaviour.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 361–371.     

Contrasting levels of genetic diversity between the historically rare orchid Cypripedium japonicum and the historically common orchid Cypripedium macranthos in South Korea

Many terrestrial orchids are historically rare and occur in small, spatially isolated populations. Theory predicts that such species will harbour low levels of genetic variation within populations and will exhibit a high degree of population genetic divergence, primarily as a result of genetic drift. If the origin of the present‐day populations is relatively recent from the same genetically depauperate source population, a complete lack of genetic differentiation between conspecific populations is expected. If a terrestrial orchid was historically common with moderate or high levels of genetic diversity, but has experienced more recent anthropogenic disturbance as a result of over‐collection, it would still exhibit initial levels of genetic variation within populations and a low degree of genetic divergence between populations. To test these predictions, we examined the genetic diversity in six populations (N = 131) of the historically and currently rare Cypripedium japonicum and in four populations (N = 94) of the historically common but now rare C. macranthos from South Korea. Fourteen putative allozyme loci resolved from eight enzyme systems revealed no variation either within or among populations of C. japonicum, which supports the first prediction. In contrast, populations of C. macranthos harboured high levels of genetic variation (mean percentage of polymorphic loci %P = 46.7; mean expected heterozygosity H_e = 0.185) and exhibited a low degree of population genetic divergence (G_ST = 0.059), supporting the second prediction. The lack of genetic variation both within and among conspecific populations of C. japonicum may suggest that populations originated from the same genetically depauperate ancestral population. The high levels of genetic diversity maintained in populations of C. macranthos suggest that the collection‐mediated decrease in the number of individuals is still too recent for long‐term effects on genetic variation. Based on current demographic and genetic data, in situ and ex situ conservation strategies should be provided to preserve genetic variation and to ensure the long‐term survival of the two species in the Korean Peninsula. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 119–129.     

The effects of cholinergic neuromodulation on neuronal phase-response curves of modeled cortical neurons

The response of an oscillator to perturbations is described by its phase-response curve (PRC), which is related to the type of bifurcation leading from rest to tonic spiking. In a recent experimental study, we have shown that the type of PRC in cortical pyramidal neurons can be switched by cholinergic neuromodulation from type II (biphasic) to type I (monophasic). We explored how intrinsic mechanisms affected by acetylcholine influence the PRC using three different types of neuronal models: a theta neuron, single-compartment neurons and a multi-compartment neuron. In all of these models a decrease in the amount of a spike-frequency adaptation current was a necessary and sufficient condition for the shape of the PRC to change from biphasic (type II) to purely positive (type I).     

Newly developed polymorphic microsatellite markers in Job's tears (Coix lacryma‐jobi L.)

A microsatellite‐enriched library of Job's tears (Coix lacryma‐jobi L. var. Ma‐yuen Stapf) was constructed using a modified biotin–streptavidin capture method. After screening, 17 polymorphic microsatellites were used for diversity analysis among 30 Job's tears accessions. The number of alleles ranged from one to five alleles per locus with an average of 2.8 alleles. Expected heterozygosity (H_E) and polymorphism information content (PIC) ranged from 0 to 0.676 and from 0 to 0.666, respectively. The newly developed microsatellite markers are expected to be very valuable tools for evaluation of genetic diversity among large germplasm collection of Job's tears present in our Korean Gene Bank.     

Spike generating dynamics and the conditions for spike-time precision in cortical neurons

Temporal precision of spiking response in cortical neurons has been a subject of intense debate. Using a canonical model of spike generation, we explore the conditions for precise and reliable spike timing in the presence of Gaussian white noise. In agreement with previous results we find that constant stimuli lead to imprecise timing, while aperiodic stimuli yield precise spike timing. Under constant stimulus the neuron is a noise perturbed oscillator, the spike times follow renewal statistics and are imprecise. Under an aperiodic stimulus sequence, the neuron acts as a threshold element; the firing times are precisely determined by the dynamics of the stimulus. We further study the dependence of spike-time precision on the input stimulus frequency and find a non-linear tuning whose width can be related to the locking modes of the neuron. We conclude that viewing the neuron as a non-linear oscillator is the key for understanding spike-time precision.     

Conditions for noise reduction and stable encoding of spatial structure by cortical neural networks

Cortical circuits have been proposed to encode information by forming stable spatially structured attractors. Experimentally in the primary somatosensory cortex of the monkey, temporally invariant stimuli lead to spatially structured activity patterns. The purpose of this work is to study a recurrent cortical neural network model with lateral inhibition and examine what effect additive random noise has on the networks' ability to form stable spatially structured representations of the stimulus pattern. We show numerically that this network performs edge enhancement and forms statistically stationary, spatially structured responses when the lateral inhibition is of moderate strength. We then derive analytical conditions on the connectivity matrix that ensure stochasticly stable encoding of the stimulus spatial structure by the network. For stimuli whose strength falls in the near linear region of the sigmoid, we are able to give explicit conditions on the eigenvalues of the connection matrix. Finally, we prove that a network with a connection matrix, where the total excitation and inhibition impinging upon a neural unit are nearly balanced, will yield stable spatial attractor responses. Received: 16 October 1998 / Accepted in revised form: 25 November 1999     

Functional redundancies, distinct localizations and interactions among three fission yeast homologs of centromere protein-B

Several members of protein families that are conserved in higher eukaryotes are known to play a role in centromere function in the fission yeast Schizosaccharomyces pombe, including two homologs of the mammalian centromere protein CENP-B, Abp1p and Cbh1p. Here we characterize a third S. pombe CENP-B homolog, Cbh2p (CENP-B homolog 2). cbh2Delta strains exhibited a modest elevation in minichromosome loss, similar to cbh1Delta or abp1Delta strains. cbh2Delta cbh1Delta strains showed little difference in growth or minichromosome loss rate when compared to single deletion strains. In contrast, cbh2Delta abp1Delta strains displayed dramatic morphological and chromosome segregation defects, as well as enhancement of the slow-growth phenotype of abp1Delta strains, indicating partial functional redundancy between these proteins. Both cbh2Delta abp1Delta and cbh1Delta abp1Delta strains also showed strongly enhanced sensitivity to a microtubule-destabilizing drug, consistent with a mitotic function for these proteins. Cbh2p was localized to the central core and core-associated repeat regions of centromeric heterochromatin, but not at several other centromeric and arm locations tested. Thus, like its mammalian counterpart, Cbh2p appeared to be localized exclusively to a portion of centromeric heterochromatin. In contrast, Abp1p was detected in both centromeric heterochromatin and in chromatin at two of three replication origins tested. Cbh2p and Abp1p homodimerized in the budding yeast two-hybrid assay, but did not interact with each other. These results suggest that indirect cooperation between different CENP-B-like DNA binding proteins with partially overlapping chromatin distributions helps to establish a functional centromere.     

Turning On and Off with Excitation: The Role of Spike-Timing Asynchrony and Synchrony in Sustained Neural Activity

Delay-related sustained activity in the prefrontal cortex of primates, a neurological analogue of working memory, has been proposed to arise from synaptic interactions in local cortical circuits. The implication is that memories are coded by spatially localized foci of sustained activity. We investigate the mechanisms by which sustained foci are initiated, maintained, and extinguished by excitation in networks of Hodgkin-Huxley neurons coupled with biophysical spatially structured synaptic connections. For networks with a balance between excitation and inhibition, a localized transient stimulus robustly initiates a localized focus of activity. The activity is then maintained by recurrent excitatory AMPA-like synapses. We find that to maintain the focus, the firing must be asynchronous. Consequently, inducing transient synchrony through an excitatory stimulus extinguishes the sustained activity. Such a monosynaptic excitatory turn-off mechanism is compatible with the working memory being wiped clean by an efferent copy of the motor command. The activity that codes working memories may be structured so that the motor command is both the read-out and a direct clearing signal. We show examples of data that is compatible with our theory.