Multimodal medullary neurons and correlational linkages of the respiratory
network. J. Neurophysiol. 82: 188-201, 1999. This study addresses the hypot
hesis that multiple sensory systems, each capable of reflexly altering brea
thing, jointly influence neurons of the brain stem respiratory network. Car
otid chemoreceptors, baroreceptors, and foot pad nociceptors were stimulate
d sequentially in 33 Dial-urethan-anesthetized or decerebrate vagotomized a
dult cats. Neuronal impulses were monitored with microelectrode arrays in t
he rostral and caudal ventral respiratory group (VRG), nucleus tractus soli
tarius (NTS), and n. raphe obscurus. Efferent phrenic nerve activity was re
corded. Spike trains of 889 neurons were analyzed with cycle-triggered hist
ograms and tested for respiratory-modulated firing rates. Responses to stim
ulus protocols were assessed with peristimulus time and cumulative sum hist
ograms. Cross-correlation analysis was used to test for nonrandom temporal
relationships between spike trains. Spike-triggered averages of efferent ph
renic activity and antidromic stimulation methods provided evidence for fun
ctional associations of bulbar neurons with phrenic motoneurons. Spike trai
n cross-correlograms were calculated for 6,471 pairs of neurons. Significan
t correlogram features were detected for 425 pairs, including 189 primary c
entral peaks or troughs, 156 offset peaks or troughs, and 80 pairs with mul
tiple peaks and troughs. The results provide evidence that correlational me
dullary assemblies include neurons with overlapping memberships in groups r
esponsive to different sets of sensory modalities. The data suggest and sup
port several hypotheses concerning cooperative relationships that modulate
the respiratory motor pattern. I) Neurons responsive to a single tested mod
ality promote or Limit changes in firing rate of multimodal target neurons.
2) Multimodal neurons contribute to changes in firing rate of neurons resp
onsive to a single tested modality. 3) Multimodal neurons may promote respo
nses during stimulation of one modality and "limit" changes in firing rates
during stimulation of another sensory modality. 4) Caudal VRG inspiratory
neurons have inhibitory connections that provide negative feedback regulati
on of inspiratory drive and phase duration.