Nervous System and

Behavior
Physiological Mechanisms:
the “How”
How Question
 Organised behaviour is a result of sensory and
motor integration – nervous system (NS)
implications
 Has behavioural needs shaped the anatomy and
organiszation of NS?
 Are there specific centres and pathways for the
control of particular behaviours?
 Are adaptive behaviours “hard-wired”
(Darwinian)?
 How is important information sifted (filtered)
from “noise”?
 Behaviour is the
tool with which
animal moves in
an organised and
directed way,
and manipulates
objects in its
environment to
suit itself.

 This chapter deals

with accounts of
behaviour in
terms of
physiological
mechanisms
(“nuts and bolts”
explanation –
HOW?)
 (Fig 3.1 & 3.2,
Barnard)
 Multicellular animals have developed complex
systems of cells
 These cells detect, transmit, integrate, store
information supplied by an animals internal and
external environment for decision making
processes:
 Sensory cells – detects changes in environment
Nerve cells – transmit and integrate information (fast)

 Chemical messengers – transmit information around the body

(slower than NS)
 Muscle cells – translate information into action
 The scope and sophistication of behaviour in
the animal kingdom is linked to the evolution
of “neural complexity”

 Vertebrate nervous systems however, are not

always more complex than those of
invertebrates
Self Reading:
 Neural anatomy and physiology
Nerve cell structure
Nerve cell function
Communication between neurones
What are the properties of the NS

that makes complex behaviour

possible?
 Evolutionary Trends in Nervous
Systems and Behaviour
As we move from unicellular organisms to
vertebrates the NS changes in 2 ways:

Greater differentiation – neurones

specialised to serve different functions

Greater centralization – neurones are

more ordered forming nerve tracts and
integrative centres (ganglia & brain)
 Phylogenetic Variation Across Nervous
Systems
 Simple Nervous Systems

 Radially Symmetrical Nervous Systems

 Bilaterally Symmetrical Nervous Systems

 Higher Invertebrates and Chordates

“Increased complexity and longer distances between parts of

the organism necessitates a more efficient form of
communication”
Simple Nervous Systems (Cnidaria)
In unicellular organisms (without NS) there
is spatial differentiation between sensory
and motor function – wave like passage of
response from one point of cell to another –
short distance

Nerve net – earliest specialised neural

organisation for conducting message between
cells – random arrangement of nerve fibres
(sea anemones, jelly fish & hydra)
Synapses are non polarised and conduct
impulse in all directions
Animals can perform simple behaviours but
no centralised processing of
information
 Radially Symmetrical Nervous Systems (Echinodermata)

 Radially symmetrical nervous system with

a circular nerve ring – starfish, brittle
stars and sea urchins (Fig. 7.6)
 Coordinated movements controlled by
impulse that travel through sensory
neurones and interneurones to the central
ring to motor neurones
 There is coordination resulting in
locomotion
 Animals with this type of nervous system
also evolved sensory receptors – contact,
taste and chemical

“Sensing the environment and coordinated responses resulted in

greater degree of flexibility and diversity of behaviours as
compared to simple systems but are still sessile or drifting”
 Bilaterally Symmetrical Nervous
Systems
 Animals here have directional movement – higher
invertebrates and chordates
 Development of cephalisation – head region where
sense organs are concentrated
 Refinement and diversification of sense organs
 Ventral surface specialised for movement
 Motor control centralised at different areas of the body –
anterior ganglia (flatworms), earthworms (single
ventral nerve cord that runs the length of the body) –
for burrowing (Fig. 7.7)
 Arthropods have larger ganglia and better sense organs
 Arthropod
Flatworm Earthworm
 In vertebrates there is further concentration and enlargement
of the nervous system – further cephalisation and
development of interconnections between nerve cells

 Vertebrate behaviour is tied to its nervous system:

1. Greater complexity and flexibility of behaviour
2. Faster responses than invertebrates due to nervous system
structure and morphology of neurones
3. Greater capacity for information storage

 Spinal cord has changed little but brain size and complexity
has changed dramatically throughout vertebrate evolution
(ratio of brain to spinal cord: fish, 1:1; humans, 55:1)
(Fig. 7.8, Table 7.1)
 Sensory Receptors (Self Reading)
 Sensation – process of transducing stimuli (sound, light, heat,
mechanical or molecules) into action potentials
 Stimuli may be external or internal to the animal – affect
sensory receptors
 Sensory processing begins in the specialised membranes of
receptor cells – affects conformational change in receptor
protein –affects membrane permeability to ion(s) – polarity of
receptor cell changes – if change is great enough – action
potential is generated
 Chemoreceptors,
 Mechanoreceptors,
 Electroreceptors,
 Thermoreceptors,
 Photoreceptors
Paramecium (unicellular invertebrate) is covered in motile cilia –
for helical propulsion in water
When it collides with an obstacle, anterior mechanoreceptors are
stimulated and cilia beat in reverse and organism move off in different
direction (Fig 3.3a, Barnard). If hit from rear, posterior
mechanoreceptors triggers forward thrust
 How do the
mechanoreceptors
communicate in absence of
nerve fibre?
Due to changes in electrical
potential of the cell
membrane which is picked up
by each cilia – entire
organism acts like a nerve
cell!
(See Fig 3.3b, Barnard)

 The control linking the

receptor to effector is
unrefined
 NS and behaviour in Invertebrates
In multicellular invertebrates the link between body parts
is taken over by axons having dendrites and synapses
Nerve Nets
 In Cnidarians (sea anemone, jelly
fish, etc) and echinoderms
(starfish, urchin, etc) (See Fig. 3.4a
& b) – they have diffused nets

 Lengthening and thickening of

axons – fast action – withdrawal
action of sea anemone (Actinaria
and Metridium)

 Behavior is stereotyped –
exhibits reflex (simple
nervous system)
Nerve tracts and Centralization  Nerve cord extends down the
 In platyhelminthes the tracts are body (Fig 3.4c, Barnard) from
more pronounced and NS shows anterior ganglion
trend towards centralization (Fig  Nerve fibres extend from the cord
3.4c) – recognisable CNS and to all regions of the body in a
cephalisation (concentration of nerve network arrangement –
tissue in the head region into an peripheral ns ( in most
anterior ganglion or simple brain) invertebrates and all vertebrates)
– simplest in flatworms
 Peripeheral nerve contain the
sensory cells and the CNS has the
motor nerve cells
 Sensory cells in Planaria respond to
touch, temp, chemicals, light
 Nerve cord allows for much rapid
transmission – increase in speed
and variety of behavioral responses
to different environmental stimuli
 The differentiation and
centralisation of NS in
flatworms allows for a degree
in learning ability – which way
to turn in a T-maze and to
avoid noxious mechanical
stimulus, assessment during
mate choice
In Advanced NS
 Sensory receptor – cell or group of
cells
 Afferent or sensory neuron – carrying
impulse from sensor
 Efferent or motor neuron – carry
impulse to effector
 Internuncial neuron or interneuron
linking sensory and motor neuron
 Effector organ (performs the motor
task)
 Nerve cords and Ganglia
In higher invertebrates like annelids (metamerically
segmented), arthropods and molluscs (non segmented) the
NS is differentiated into ganglia linked by nerve cords
Increasing centralisation – neural
switchboard:
 Afferent fibres (sensory receptors) connect to
interneurons and to motor neurons
 Depending on input, different motor neurons
are brought into play
 In leeches ganglia has 400 cells; in Aplysia a
mollusc has 1500 cells
Function of ganglia:
 regulation of local reflex arc – local control

 Long range coordinated control via long

interneurons along nerve cords facilitating
coordinated operations of different body
parts
 A number of behavioral advances are associated with
these developments
 Elaboration of appendages and musculature

Emergence of fluid filled body (coelom)

 allows for subtle movements and complex
manipulative tasks eg. In the web building
 Elaborate courtship songs and ornamented nest
construction of some insect species
 Stimulus discrimination and learning but learning is
short lived (small capacity of the ganglia) just like the
short live span of species
 Among invertebrates there is trend towards
enlargement of brain – amalgamation of
somatic ganglia

 Somatic ganglia still retain considerable

independence of control

 If cerebral ganglia is removed earthworms still

can crawl, feed, copulate but are hyperactive

 Nereids are able to learn certain tasks even after

removal of cerebral ganglia from CNS
NS and Behaviour in Vertebrates
 NS develops from dorsal tissues and as a tube rather than as a solid
structure
 Traces of ancestral segmented patterns are also present – distribution of
sensory and motor neurons
 Centralisation, cephalisation and functional differentiation reaches its peak
in vertebrates
 Structure and function of CNS (See Fig 3.6, Barnard)

Brain and spinal cord

 Brain has 3 regions and can be further sub-divided: forebrain
prosencephalon), midbrain (mesencephalon), hindbrain (rhombecephalon)
 Evolutionary trends in the vertebrate brain
2 main evolutionary trends:
 First, elaboration of
the midbrain (shown
in fish) – optic tectum
thickens and stratified –
a integration center for
information from other
parts of brain
 Second, elaboration of
the cerebral
hemisphere (shown in
mammals) shown in the
forebrain – major
association centres (See
Fig 3.7a & b, Barnard)
 Forebrain of lower
invertebrates remain in
the form of hippocampus
but the neo-cortex in
humans extends to the
whole of the brain
 Plot of brain and body
size (See Fig 3.7c,
Barnard). The
discontinuity is a result of
the elaboration of the
forebrain cortex (neo-
cortex) in higher
vertebrates
 In lower vertebrates
brain is <0.1% of body
mass, in birds and
mammals > 0.5%, in
humans it is 2.1% (See
Fig. 3.8, Barnard)
 Neo-cortex has many visible
divisions which reflect
different functional areas (See
Fig. 3.8a, Barnard)
 In advanced mammals
voluntary motor control are
in front of somatic sensory
functions and separated by
deep fissures
 Part of the motor cortex
communicates with the spinal
cord via the pyrimidal tract
(see Fig. 3.7b, Barnard)
 The 2 halves of the cortex are
connected by the corpus
collosum
 Severence of the corpus
collosum results in loss of
speech control and verbal
comprehension
 Further motor control is at
the corpus striatum
(subcortical); in birds it is
important for sterotype
bahaviour
 The limbic system
(subcortical) – hippocampus,
cingulate gyrus, septum and
amygdala – control arousal,
learning, agnostic behaviour
and decision making (See
Fig. 3.8b, Barnard)
 The thalamus – relay
information from retina,
ear, cerebellum, and tectum
and function in appreciation
of temp, pain and pleasure;
production of hormones in
hypothalamus – emotional
arousal, sleep, feeding,
aggression, osmoreceptors
(drinking, orientation
behaviour)
NS and the Adaptive Organisation of
Behaviour
 There is association between
organisation of NS and the range and
complexity of behaviour of different
taxonomic groups

 Behaviour results from coordinated

neural control of the effector system

 Variation in organisation reflects adaptive

specialisation between and within species
 Does gross anatomy of CNS reflect
different adaptive behaviour patterns?

 Are adaptive behaviours “hard wired”

into nervous systems – do they have
neural circuits dedicated to their control?

 What can we infer about behaviour

specialisations from anatomy of nervous
system from evolutionary trends moving
towards centralisation and
cephalisation?
 Comparative studies of
Invertebrates
Lifestyle
 Early development are nerve nets –
cnidarians – rapid through conduction –
quick response to noxious stimuli.

 This fast through conduction is also in

annelids and arthropods in the form of
large axons – giant fibres.
The annelids and arthropods have wide
range of adaptive behavioural
specializations. Are these reflected in the
anaotomical arrangement of the fibres?

There is difference in the oligochaetes

(earthworms) and polychaetes
(ragworms) – anatomy and conduction of
giant fibres with respect to their lifestyle
 In Lumbricus (burrowing
oligochaete) (See Fig 3.9a)
the primary function of giant
fibres is for fast contraction
of longitudinal muscles and
forward and reversal
withdrawal in response to
mechanical stimuli –
emergency responses in
burrowing organism

 The giant fibres makes up

10% of cross section area of
the nerve cord
 Among polychaetes the giant fibres make up 25-70% of the cross section of nerve cord (Fig.
3.9b, Barnard)

 Insedentary fan worms (Myxicola, Sabella, Branchiomma) the function is for withdrawing the
feeding mechanisms into burrow
Errant polychaete (Neries) – are active surface
predators:
 have complex eyes and
 sensory tentacles
 large and well differentiated brain
 greater locomotory actions (withdrawal and
creeping),
 side-side swimming
 rotary motion of the parapodia
 The giant fibres in Nereis are
same as in Lumbricus but there
is thickening of the lateral
fibres (Fig 3.9c, Barnard) –
have extensive connections and
closer association with motor
axons

 The 3 central fibres (median

and paramedian) have control
over parapodia while control
of longitudinal muscles is by
lateral fibres

 The division of labour has

allowed for rapid but diverse
locomotory control which is
important to a mobile predator
 • The brain of anelids and arthropods share broadly similar
glomeruli
• The relative sizes of glomeruli is related to differences in
behaviour
Hanstrom (1928) compared
glomeruli between actively
hunting lycosids (wolf
spiders) and web spinning
agelenids (Fig. 3.10):

Lycosids have extensively

developed optic centres and
corpora pedunculata which
are glomeruli for sensory
association and visual
memory

Agelenids have larger

central body which is
associated with integration
of notably preprogrammed
behaviour
 Their anatomy thus reflects the different
lifestyles of the two groups and their
different demands on sensory integration
and motor skill

 A comparison across the insects shows a

general association between the size and
structure of the corpora pedunculata and
behaviour particularly to
 Social organisation and
 Spatial complexity of foraging behaviour
 Comparative Studies in Vertebrates
Lifestyles
 Broad behavioural characteristics is correlated with brain – the
relative sizes of the different parts of the brain (see Fig 3.8,
Barnard)
 As a group, otters have a range of foraging
skills and manipulating food with their fore
paw, but these skills vary with species (See
Fig 3.11)

 Sea otter (Enhydra lutris) – clawless – breaks shellfish

against stone placed on its chest

 Clawed relatives (Lutra canadensis – river otter;

Pteroneura braziliensis – South American giant otter)
use their fore paws in a much less specialised way –
more emphasis on sensory information face and vibrissae
 Radinsky (1968) showed differences in the cortex of the two
groups (See Fig 3.11, Barnard) – forelimb projection in cortex
for handlers larger while sensory projections in cortex larger
for face and vibrissae information
 Bird Songs range from simple notes to complexity
and melody.
 Song control system is the discrete nuclei in the
forebrain that is projected to the syrinx (vocal organ)

 The nuclei are of 2 groups (See Fig 3.12a, Barnard):

 Higher vocal centre (HVC) and robustus
archistrialis (RA) – role in song production
 Area X and lateral magnocellular nucleus (l-
MAN) – song acquisition
DeVoogd et al. (1993)
compared HVC and
Area X in 45 songbird
that differed in song
complexity – repertoire
size and number of
syllable per song by
using DNA phylogeny
• There was positive
correlation between size
of song repertoire and
volume of HVC (Fig.
3.12b)
• Species with larger
repertoires had larger
nuclei associated with
control of song
production
Sex Differences in Brain Structure
NS differs between males and females in
relation to
 reproductive roles,
 ecology
 many differences are also driven by hormones
mediated by sex differences of hormone receptors in
the brain
 In human there are 3 differences in structural
architecture of the brains of males and females
resulting in different sexual behaviour
49
First
dimorphism in size of nucleus INAH-3 which is 2
to 3X larger in males and packed with androgen
sensitive cells – male typical sexual behaviour

High levels of androgens in women – male like assertive

sexual behaviour, small breasts, low vocal pitch,
hirsuteness (hairy)
Controversially INAH-3 has been linked to male
homosexuality (Levay, 1991 – studied males who died of
AIDS and men who did not die of AIDS)
INAH-3 nucleus in homosexuals smaller than in
heterosexual males and roughly the same size as that of
women
Second
connection between the cerebral hemispheres
(the corpus collossum, anterior commisurre,
and nerve fibre bundles) are larger in women
 Brings about greater empathy and emotional
sensitivity.
 Women have greater connection between the 2
halves of the thalamus which is important for the
relay of sensory information to cortex
Third
Ageing – men lose more brain tissue as
they age and earlier in life
 Tissue loss is in frontal and temporal lobe in
men; hippocampus and parietal area for
women
 Accounts for personality and behaviour
changes with age
 Increased irritability for men and reduced
memory and visual skills for women
Ecology and Sexual Dimorphism in Brain
Structure (long term selection pressure)
 Sexes differ in general features and brain anatomy but
specific differences can arise due to ecological selection
pressure – eg. spatial memory and navigational skills
(greater in males)
 Males have larger home range or territories – greater spatial
learning tasks. Do males have greater development of the
brain related to spatial memory, ie. the hippocampus
(spatial awareness and navigation)?
 Studies show that males do have larger hippocampus (for
birds and mammals) – but there are exceptions, however!
54
 Sherry et al. (1993) - brown headed cowbirds
(Molothrus ater) – brood parasites
 Females search for host nests and lay single egg in
each nest – up to 40 eggs in each breeding season.
Males play no part.
 Sherry also compared with closely related non
parasitic birds (control species – red winged
blackbird and the grackle)
 Sex differences in brain structures do not always
reflect differences in associated behaviour.
 Sherry et al. (1993) hypothesized that spatial abilities of females
(locate and return to host nest) would show larger hippocampus than
males – females do have larger hippocampi than males (Fig. 3.13)
 Clayton et al. (1997) however, stated that sex
difference in size of cowbirds hippocampus (female
and male) was related to breeding season when birds
are actively looking for host (cyclical or seasonal)

 The size of forebrain nuclei (controlling song) in

canaries also show cyclical variation with season and
thus song production

 This suggest adaptive plasticity in brain

development related to the environment
Brain Development and Experience
 Some species of birds store food for later retreival –
spatial memory required to retrieve the items
 Black capped chikadees (Parus atricapillus) scatter-hoard seeds
and insects in bark and moss clumps over a wide area
 Items are hidden individually and each location used only once
and can retrieve the food items even after a month! (Hitchcock &
Sherry, 1990)
 Birds remember specific sites – the Clarks’s nutcracker
(Nucifraga columbiana) can hide 9000 items and can find them 9
months later – birds really do remember specific sites - not
using approximates of visual or odour cues to detect food about
the area to search for food
 Birds not only can remember what items they hide in different
places but also can time the recovery of the items according to
how fast they decay – episodic memory
 The region of the brain involved in the episodic
memory is the hippocampus – so food hoarding
species would have larger hippocampus than closely
related species that that do not – this is the case!

Size of hippocampus appears to be affected by

the hoarding experience of individual birds
Clayton & Krebs (1994)
experiment on marsh tits
(Parus palustris) –
divided birds into 2
groups – first group
allowed to cache
sunflower seeds and
second not able to

They then examine

brain of both groups –
second group had smaller
hippocampi (Fig 3.14 b) –
experience of hoarding
had profound effect on
hippocampus
development
 Macguire et al. (2000)
used MRI to study
the spatial learning
in taxi drivers

 Taxi drivers who had

completed training
had larger posterior
hippocampus as
compared to controls
and increased with
the length of time the
drivers had been
operating (Fig 3.14c)
 Is Adaptive Behaviour “hard wired”?

 In previous section – adaptive behaviour patterns can

be linked to particular areas of the brain
 Size of these areas reflect the relative importance of
the behaviour in different species and individuals
 Is there a particular component of the ns as the
mechanism controlling a behaviour?
 Do adaptive behaviours have neural circuits?
 Answer is YES and it depends on the definition and
complexity and on the nature of information
processing within the ns
 Local versus distributed processing
 Before 1990 – behaviour was thought to be
mapped to simple neuronal circuits – local and self
contained
 Now it is thought as neural networks (different
ways and different times) and distributed
processing system (at immediate site and away –
Connectionist view – even simple tasks may
require 100 million cells or more (John et al.,
1986)
Reflexes
 A simple form of behaviour – an automatic, quick,
stereotype unit of behaviour in response to a stimulus
(internal and external)
 Examine the relationship between behaviour and the
functioning of specific neural circuits – eg., knee-jerk
response and limb withdrawal from a painful stimulus
(Reflex arc, Fig 3.15)
 Limb withdrawal reflex involves
 Simultaneously contract flexor muscles and relax the
antagonistic extensor muscles so that limb can be pulled
towards body
 1st – flexion reflex – as affected limb is flexed, has to use
other muscles to steady itself (cross-extension reflexes) –
in combination the 2 allow for emergency actions and
locomotion
 Stretch reflexes – to grade flexion and extension in
antagonistic muscles so that limb is controlled in stages
and not in one violent action
 There is no elaborate CNS in invertebrates but sensory and effector organs still
communicate through the central nerve cord and ganglia, eg. in the gill withdrawal
response of sea hare Aplysia (a mollusc)
 The gills are retracted into the mantle cavity in response to weak mechanical
stimulation of the siphon
 The reflex arc involes – excitatory and inhibitory interneurons relaying sensoty
information from siphon to motor neurons in the abdominal ganglia which causes
withdrawal of gill (Fig 3.16)