Inhibition and excitation peak with the activity, we first examined the origin of the total neuron conductance during the depolarizing waves of scratch episodes. Furthermore, spike periodic high-conductance state reported in activity is driven by depolarizing synaptic transients rather than pacemaker properties.
These motoneurons during scratch episodes The findings show that balanced excitation and inhibition and irregular firing are fundamental motifs periodic nerve activity recorded in parallel from in certain spinal network functions. This reciprocal arrangement Department of Neuroscience and Pharmacology, Panum T generating motor circuits in the spinal cord proposes that excitatory interneu- rons in each half-center drive agonist motoneu- predicts half-center neurons to be excited and inhibited in alternation during rhythmic network activity.
E-mail: motoneurons and interneurons in the antagonist to operate at low intensity of synaptic activity j. Inhibition and exci- tation covary in motoneu- rons during scratching. F Relation between normal- ized increase in inhibitory F 1. Blue arrows represent Inhibition leading single trials, and red arrows the trial average, for a single 0 cell G , weighted average for 0 0.
The membrane potential strychnine 0. Reduced local inhibition 0. With the simplifying Balanced increase in excitation and inhibi- 3D.
Finally, regular firing with highly correlated assumptions proposed by Borg-Graham et al. We therefore of inhibition and excitation 0.
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First, the reg- low-amplitude depolarizing waves aided by Fig. In the cell illustrated and ular firing in motoneurons, induced by a steady 1. These conditions also reduced the also Ge and Gi peaked with each depolarizing activity purple in Fig. Suc- membrane potential to a minimum.
Polar plots of after-hyperpolarizations In contrast, firing from network neurons located far enough below Downloaded from www. Spike-triggered previous theoretical and experimental findings depolarizing waves would contribute to the in- averaging revealed that APs during irregular in other systems. The balanced state hampers creased Gi 16, This was not a major factor, firing are preceded by a brief depolarizing tran- regular firing by increasing conductance and pro- however, because high-conductance states and sient arising from a flat average voltage trajec- motes irregular firing by increasing fluctuations in the coherence between Ge and Gi remained in tory Fig.
S3 in membrane potential 21, 23— Thus, sponse properties is severely obstructed by a duced by using recording electrodes containing irregular firing is induced by depolarizing conductance increase of the magnitude observed QX and CsCl Fig. The re- synaptic transients in the high-conductance state during the depolarizing waves of scratch epi- maining explanation is that there is strong during network activity. A possible source of sodes At the same time, however, the 2- to inhibitory synaptic activity during the depolariz- these transients is a high incidence of uncorre- 5-fold increase in conductance that brings the ing phase of the scratch.
We tested the sensitivity of irregular potentials is associated with a parallel increase tion was reduced pharmacologically Fig. During scratch in tegrated power spectrum in the Hz band input to the recorded motoneuron was reduced control conditions Fig. The broad spectral content asso- ciated with the rhythmic activity is fully in line with predictions for a state of intense and bal- anced inhibitory and excitatory synaptic activity A B 16, 19, 21, 24 and orders of magnitude higher than expected for channel noise The spinal network studied here shares sev- eral properties with the balanced state in mathe- matical models of large-scale random networks of inhibitory and excitatory neurons 28, Depolarizing waves enhanced by local reduction in inhibition.
B Single fig. This suggests that the scratch network wave highlighted. Regular firing by intrinsic properties at rest, irregular firing by synaptic transients during network activity. A Regular firing in moto- neuron at rest purple replaced by irregular firing during network activ- ity blue induced at the onset of mechanical stimulus vertical arrow.
B Spikes triggered by pacemaker potential at rest and by depolarizing transient during network activity. C to E Increasingly regular firing left and correlated ISIs right during scratch in control C , reduced inhibition D , and reduced inhibition and excitation E. Aligned recordings Downloaded from www. Highlighted depolar- izing wave middle.
The intensity of synaptic fluctuations Membrane potential Upper trace Sample intracellular recording obtained with —2. Middle trace Gtotal obtained as in Fig. Lower trace The integrated to 25 mV Hz band power spectrum of the subthreshold membrane potential. The high-conductance state, how- with functional motor networks in the spinal out repeating the spike patterns of individual ever, sacrifices the temporal dynamics offered cord. The straightforward functional correlate neurons. This is in agreement with the chaotic by models based on weakly coupled neurons and absence of anesthetics and other drugs nature of the balanced state in mathematically with oscillatory intrinsic properties 6, 7.
It makes our experimental model appealing in modeled networks in which the activity of remains to be seen whether high-conductance the search for computational advantages that individual neurons is stochastic and highly states occur throughout the scratch network or balanced inhibition and excitation may pro- sensitive to initial conditions Adopting a only in the motoneurons and large interneu- vide in large-scale neural networks in general. Sherrington, The Integrative Action of the Nervous robustness of the spinal scratch generator, its of inhibitory and excitatory synaptic activ- System Cambridge Univ.
Press, Cambridge, Graham Brown, Proc. London B. Grillner, in Handbook of Physiology, Sect. Figueres, C. Kollmuss, A. Meinshausen, M. Pacala, S. Ramanathan, V.
Rogelj, J. Brown, M. Busby, J. Campbell, K. Dasgupta, S.
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Michael, J. Voinov, A. Weitzman, M. Alsop, Z. Campbell-Lendrum, D. Costello, A. Epstein, P. Haines, A. Lim, V. McMichael, A. Patz, J. Begley, S. Jones, D. Peterson, T. Preprint here. Revkin, A. The emissions growth rate since was greater than for the most fossil-fuel intensive of the Intergovernmental Panel on Climate Change emissions scenarios developed in the late s. The growth rate in emissions is strongest in rapidly developing economies, particularly China. The results have implications for global equity. The time series are constructed using direct measurements wherever possible and reconstructions based on models and proxies at earlier times.
These time series are compared with the climate record for the period to The solar records are scaled such that statistically the solar contribution to climate is as large as possible in this period. Under this assumption we repeat the comparison but now including the period This comparison shows without requiring any recourse to modeling that since roughly the solar influence on climate through the channels considered here cannot have been dominant. Current knowledge cannot rule out a return to such conditions in response to continued greenhouse gas emissions. Moreover, a threshold triggering many meters of sea-level rise could be crossed well before the end of this century, particularly given that high levels of anthropogenic soot may hasten future ice-sheet melting …, the Antarctic could warm much more than , years ago [when sea levels were m higher] … and future warming will continue for decades and persist for centuries even after the forcing [the gas levels causing climate change] is stabilized.
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Continued growth of greenhouse gas emissions and associated global warming could well promote SLR of 1m-3m in this century, and unexpectedly rapid breakup of the Greenland and West Antarctic ice sheets might produce a 5m SLR. In this paper, we have assessed the consequences of continued SLR for 84 developing countries. Geographic Information System GIS software has been used to overlay the best available, spatially-disaggregated global data on critical impact elements land, population, agriculture, urban extent, wetlands, and GDP with the inundation zones projected for m SLR.
Our results reveal that hundreds of millions of people in the developing world are likely to be displaced by SLR within this century; and accompanying economic and ecological damage will be severe for many.