Respiratory Physiology & Neurobiology 154 (2006) 61–72
Modulation of aerial respiratory behaviour in a pond snail
Ken Lukowiak
∗
, Kara Martens, Mike Orr, Kashif Parvez,
David Rosenegger, Susan Sangha
Hotchkiss Brain Institute, Faculty of Medicine, University of Calgary, Calgary, Alberta, Canada, T2N 4N1
Accepted 10 February 2006
Abstract
Aerial respiratory in Lymnaea is driven by a three-neuron CPG whose sufficiency and necessity has been directly demonstrated.
While this CPG is ‘hard-wired’ it displays a tremendous amount of plasticity. That is, it is possible by employing specific training
procedures to alter how it functions in a specific hypoxic environment. Thus, it is possible to study directly the causal mechanisms
of long-term memory formation, forgetting, and modulation of the memory at a single cell level. Thus, it is possible to use a
relatively simple three-neuron CPG to study not only important questions concerning regulation of important homeostatic
mechanisms but to also use it to study how learning and non-declarative memory are mediated at a cellular level.
© 2006 Elsevier B.V. All rights reserved.
Keywords: Respiratory CPG; Learning; Memory; Lymnaea
1. Introductory remarks
As physiologists in one-way or another our research
is concerned with ‘the where’, the ‘why’ and ‘the how’
of homeostasis and quite possibly the neural control
of respiration is the quintessential homeostatic system.
Our laboratory has focused on the neuronal mecha-
nisms underlying control of two molluscan (Aplysia
and Lymnaea) respiratory systems. However, our pri-
mary reason for using these model systems was not to
This paper is part of a special issue entitled: “Frontiers in Com-
parative Physiology II: Respiratory Rhythm, Pattern and Responses
to Environmental Change”, guest edited by W.K. Milsom, F.L. Pow-
ell and G.S. Mitchell.
∗
Corresponding author. Tel.: +1 403 220 6872;
fax: +1 403 283 2700.
E-mail address: lukowiak@ucalgary.ca (K. Lukowiak).
study respiratory behaviour per se; but rather to use
these systems to study the neuronal mechanisms of
learning, memory, and forgetting. We will concentrate
this review on key aspects of aerial respiration in Lym-
naea and show how this behaviour can be modified by
experience, the environment, and ageing.
Homœstasis is defined as in the following manner:
“to designate stability of the organism” and according
to the OED was first used in the English language by
Walter Canon in 1926. Unsurprisingly then the word
may imply to some the necessity of constrained plas-
ticity in the neural system that controls a specific home-
ostatic behaviour; however, we feel that the ability to
maintain homeostasis can only come about because
there is plasticity within the neural control system. Let
us explain what we mean by this. The ‘hard-wired’
neural system that controls aerial respiration in Lym-
1569-9048/$ – see front matter © 2006 Elsevier B.V. All rights reserved.
doi:10.1016/j.resp.2006.02.009