Anim. Behav .
, 1978, 26,969-981
THE INSECT NERVOUS SYSTEM AND INSECT BEHAVIOUR*
BY FRANZ HUBER
Max-Planck-Institut fur Verhaltensphysiologie, Seewiesen, Germany
A Well-Remembered Conversation
My personal contacts with Niko Tinbergen go
back to March, 1952, when there was a meeting
of ethologists in Westphalia . Colleagues from
England, Holland, Scandinavia, Switzerland and
Germany all converged on the castle of Duke
Rhomberg in Buldern, where Lorenz's group
was working. At that time I was still a zoology
student in Munich, with Karl von Frisch and
Werner Jacobs, and I was given the chance to
speak in Buldem about my first results with
local brain lesions in crickets and grasshoppers
and their effects on singing behaviour . One
outcome of this talk was a walk with Niko
Tinbergen in the castle park ; he wanted to hear
all the details of my approach and methods . In
our conversation he convinced me that the future
of ethological research would emphasize experi-
mental analysis, as well as observation of the
natural behaviour. Ethology, he said, would
need young researchers familiar both with the
literature and methods of ethology and with
those of physiology ; neurobiology in particular .
Behavioural physiologists should be trained to
formulate questions and design experiments that
are biologically relevant and appropriate to the
behaviour of the animal under study . It must be
the formulation of the problem that determines
the experimental procedures, and not the other
way around ; the temptation to ask only questions
answerable by a favourite method must be
resisted! Niko Tinbergen emphasized that the
etho-physiologists this new kind of training
would produce should know the life history of
their chosen animals and, if possible, the
ecological conditions limiting their ranges .
With this background, they could avoid experi-
mental conditions that disturb behaviour and
be in a better position to account for normal
behaviour. And he laid particular weight on a
proposition that K. Lorenz and E . v. Hoist had
also often put forward : the behavioural physi-
ologist ought to begin his study with the intact
organism. Even when it is the nervous system
that interests him, he should proceed in such a
way that his results illuminate the neural basis
of behaviour. Today we know that these ideals,
*The Niko Tinbergen Lecture, 1977 .
969
justified as they are, can be realized only within
limits. But as a description of the ultimate goal
of research, Niko Tinbergen's comments in-
fluenced me profoundly. These, together with
Kenneth Roeder's (1963) admirable approach
to analysing the neural basis of insect behaviour
had a decisive effect on the conceptual and
experimental approach to insect neuroethology
that I, and the members of my group, were to
follow.
Since then, whenever possible, our studies of
the neural basis of auditory behaviour in crickets
and grasshoppers have employed freely moving
animals with a minimally restricted radius of
activity . It proved possible to stimulate localized
regions of the small insect brain under these
conditions and to elicit singing behaviour
(Huber 1960 ; Otto 1971 ; Plate XVI, Fig. 1) .
This method has since been extended to include
continuous observation of the animal by a TV
monitor and computer processing of the res-
ponses to stimuli (Wadepuhl & Huber, in
preparation).
In much the same way, we have used im-
planted recording electrodes to study the neuro-
muscular organization of singing movements and
the more complex courtship activities, designing
the system to maximize freedom of movement
and naturalness of behaviour (Huber 1965,
1975 ; Elsner 1968 ; Kutsch 1969 ; Plate XVI Fig.
2 and Plate XVII, Fig . 3) . Even in our work on
the processing and recognition of the sounds
biologically relevant to crickets, we make every
effort to use intact moving animals for our
recordings (Plate XVII, Fig . 4). One of our aims
here is to monitor the neuronal processing of a
female engaged in phonotactic orientation, the
motivational state of which is known (Stout
1971).
I must admit that we have not yet reached this
goal . For the time being we must often rely on
`preparations', that is, fixed animals with nervous
systems partially exposed, which cannot behave
normally . Such preparations do of course have
their value . We can study the input/output
relationships of single neurons, for example,
which are enormously important to an eventual
understanding of the neural basis of behaviour .
And we can work out the interconnections of