Lecture 13

INBORN & ACQUIRED FORMS OF BEHAVIOR

PHYSIOLOGY OF HIGHER NERVOUS ACTIVITY

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 LECTURE OUTLINE

1. Definitions of lower and higher nervous

activities. Forms of behavior
2. Inborn forms of behavior. Characteristics of
unconditioned reflexes. Instincts
3. Acquired forms of behavior. Conditioned
reflexes
4. Physiological rules of conditioned reflexes
formation
5. Dynamic stereotype
6. Types of conditioned reflexes inhibition
7. Types of HNA in animals and human

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 NERVOUS ACTIVITY
LOWER NERVOUS ACTIVITY HIGHER NERVOUS ACTIVITY
• The lower nervous activity (LNA) • Higher nervous activity (HNA) provides
supplies the primitive regulation adequate individual behavior in the
of organs and systems and their environment and one’s adaptation for
integration into the whole different exchanges of external and
organism as well as elementary internal environment.
adaptive responses
• The main advantage of HNA is the
• Adaptive capabilities of the adaptive reactions of outstripping
lower nervous activity are limited; reflection, i.e. before the stimulus
they may be applied to the action.
stereotypical situations
• Higher nervous activity comprises the
• Lower nervous activity is based most complex forms of behavior that
on the inborn reactions and are acquired during individual
behavior forms. development of person.
• unconditional reflexes, • imprinting,
• instincts, • conditioned reflexes,
• biological motivations • thinking,
• emotions • cognition. 3
 1863 – “Reflexes of the brain”
Russian physiologist. Was the
first who developed, in a broad
scope and details, the
conception of the reflex
character of the higher divisions
of the CNS in his monograph
“Reflexes of the Brain”.
His discovery made the
possibility of an objective, purely
physiological analysis of the
psychic processes which were
SECHENOV regarded as the exclusive of
Ivan Mikhailovich subjective psychology

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 • Has developed Sechenov’s ideas
about the reflex principle of CNS
action
• Elaborated method of conditioned
reflexes which is the objective
experimental method for
investigation of the cerebral cortex
functions
• Elaborated the conception of
higher nervous activity.
• Demonstrated that reflex reactions
in the lower divisions of the CNS
(subcortical nuclei, brainstem and
spinal cord) were accomplished
by inborn inherited nervous
pathways, whereas nervous
connections in the cerebral cortex
PAVLOV were developed anew, in the
process of individual life of animals
Ivan Petrovich and human.
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 Inborn forms of behavior
• Unconditioned reflexes are inborn, inherited
reactions of an organism in response to internal and
external stimuli that are realized with participation
of nervous system.
• Unconditioned reflexes arise just in response to
immediate (direct) irritation of the receptive field;
their realization doesn’t require special conditions.
• Irritant which causes the specific reflex reaction is
called unconditioned stimulus.
• The principle of unconditioned reflexes:
• stimulus → response reaction → back afferentation
• If there is no stimulus, there is no response reaction!
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Peculiarities of unconditioned reflexes

1. Unconditioned reflexes are inborn, inherited

• They are realized according the strong
genetic program
• The most of them are exhibited right after birth
(for instance, the sucking reflex of mammals,
sneezing, coughing, swallowing, salivation)
• Some unconditioned reflexes (for instance,
sexual reflexes) are formed after the
appropriate development of nervous,
endocrine and other systems.

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 Peculiarities of unconditioned reflexes
2. Unconditioned reflexes are specific reflexes that
are inherent for all specimens without exceptions.
3. These reflexes are permanent. They are very
stable and are maintained during not only the single
individual life, but the whole species existence.
4. Unconditioned reflexes arise when the adequate
stimulus acts to the specific receptive field

• For example, at the natural conditions the unconditioned

reflex of salivation can be observed when the food
(adequate stimulus) acts to the mucosa of oral cavity (its
specific receptive field).

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 5. Reflex arch of unconditioned reflex
3 links = 6 components

• Nervous center has a multilevel structure

• It consists of some departments that are located at the
different levels of CNS – spinal cord, medulla, brainstem, etc.
• The highest department of nervous center is situated at the
cortex hemispheres, and is called cortical representation of
unconditioned reflex
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 Classification of unconditioned reflexes

Sechenov Pavlov Kanorsky

 food 1. Vital – ensure individual and 1. Keeping
 defense specific conservation: (retaining)
 sexual food, drinking, sleeping
 orientation regulation, defense, orientation
 paternal reflexes
 infantile 2. Role – zoosociological 2. Defense
unconditional reflexes that can be
realized only when interaction
with another individuals of the
species
3. Self-developing – research ,
opposition (freedom), imitating,
playing

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 Instincts
• Instincts are inborn form of
behavior.
• Instinct is the complex system
(sequence) of unconditioned
reflexes in response to the
irritation from the external or
internal environment
• Instincts have a chain nature and
the end of one reflex is the
beginning of the next one at that.
• The manifestation of instincts
depends upon the numerous of
internal and external factors:
• 1) specific conditions,
• 2) metabolism,
• 3) humoral status,
• 4) intellectual process.
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 Biological significance of instincts
• The strong specific uniformity of instincts provides
the maintenance of vital functions to survive
individual and species, independently from the
accidental ambient conditions

Instincts are not just the

basis of organization of
animals’ behavior, but they
are also fundamentals for
evolution to produce more
complex acquired forms of
behavior

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Acquired forms of behavior
IMPRINTING
• The simplest way of learning is
imprinting. It is observed in children
before 5 years old

• Experiments of Conrad Lorentz with

goslings

• Imprinting - imitative behavior – is the

intermediate between unconditioned
and conditioned reflexes and has the
features of both

• For instance, the reaction of following

is inherited, but to cause this reaction
the specific condition is necessary –
moving object

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 CONDITIONED REFLEX

• Is the fundamental and the most various form of

the purposeful behavior organization
• The crucial factor to produce the conditioned
reflexes and entire adaptation of an organism is
the interaction of central excitation and central
inhibition
• The manifestation of higher nervous activity
depends upon the properties of main nervous
processes which totality characterizes the
individual peculiarities of behavior.

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 Comparative characteristics of
unconditioned and conditioned reflexes
Unconditioned reflexes Conditioned reflexes
(inborn behavior forms) (acquire behavior forms)
Can be realized right after the birth Are formed during the individual
moment (sucking, swallowing, development of person
sneezing, coughing) or later (sexual
reflexes)
Specific – are inherent for all Individual, depend upon the
specimen without exception conditions of existence
Permanent – are realized during Variable – arise in the definite
entire human’s life conditions and disappear in the
absence of these conditions.
Require the adequate irritant action Are formed under the influence of
immediately to the specialized numerous of stimuli
receptors (food – taste receptors, light
– visual receptors)

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The rules of conditioned reflexes formation
1. Availability of two stimuli:
1) conditioned (sign) irritant, that carries information about
the further unconditioned irritant (light, sound, smell of
food) and
2) unconditioned stimulus, or unconditioned reinforcement
(food, pain) which elicits unconditioned reflex.

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The rules of conditioned reflexes formation
2. Temporal factor
1) Conditioned stimulus must precede unconditioned one
2) Temporal interval between conditioned and unconditioned
irritants must be short (optimum is 20-30 seconds). Duration is
in the direct dependence from strength and information
significance of irritant
3) To form conditioned reflex multiple repetitive combination is
necessary: conditioned signal – unconditioned irritant
(sometimes it’s possible to produce the conditioned reflex
after a single combination)

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The rules of conditioned reflexes formation
3. Factor of strength:
1) The strength and biological significance of conditioned irritant
must be less than the unconditioned one
2) The strength of conditioned stimulus must be predominant
against other indifferent stimuli – presence of motivation.
For example, to produce the conditioned reflex of salivation,
the experimental animal must be hungry.
3) Both conditional and unconditional irritants must have an
optimum strength – “the law of strength relations in the cortex”
4) To form conditioned reflex the normal brain activity is necessary
(consciousness, mental health)
5) Individual must be healthy somatically
Conditioned reflexes are formed on the basis of unconditioned reflexes

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Significance of temporal-spatial connection
• The physiological basis of the
conditioned reflexes is the
formation of temporal-spatial
connections in the highest
departments of CNS
• According I. P. Pavlov, it is formed
between cortical part of analyzer
and cortical representation of
unconditioned reflex)
• Temporal-spatial connection is
the totality of neurophysiological,
biochemical and ultrastructural
remodeling in the brain occurring
during the cooperative effect of
both conditioned and
unconditioned stimuli

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 Scheme of conditioned reflex formation
Conducting
I. Conditioned department
stimulus Analyzer
Cortical
(light, sound) receptors part of
Cortical analyzer
representation
of UR
Hth
After 20-30 +
seconds MOTIVATION ++
+
TEMPORAL-
SPATIAL
II. Unconditioned Receptive CONNECTION
stimulus field
(food) Center of
unconditioned
reflex

To the organ-
effector

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 Conditioned reflexes formation
Excitation caused by the indifferent (conditioned) stimulus from any
receptive fields is transmitted by afferent nerve fibers into the cortex
hemispheres where the region of excitation arises

If in 20-30 seconds the animal receives unconditioned reinforcement

(food), in the cortex new region of excitation will appear

This excited region is stronger than previous one, because the

unconditioned stimulus (food) always has a greater biological
significance for animal than the light or sound

The region of unconditioned excitation is the dominant center. As a

dominant center, it attracts excitation from the region of conditioned
excitation, and initiate reflex which it controls (salivation)

Repeated combination of conditioned stimulus and unconditioned

reinforcement facilitates impulses conduction between two centers
(principle of “beaten track”) – temporal-spatial connection is formed 21
 Common features of conditioned reflexes
• Conditioned reflex has adaptive character.
• The meaning of conditioned reflex is to produce the behavioral plasticity,
correspondence of behavior to the current conditions (time, place,
reinforcement)
• Conditioned reflexes are formed with participation of higher levels of CNS
• Conditioned reflexes are produced and eliminated during the individual
life of person.
• The most of conditioned reflexes are produced after some repetitions of
conditioned and unconditioned signals. In other words, individual
memory doesn’t recode every random coincidence of conditioned and
unconditioned irritants, just most probable of them. But if the probability to
receive the reinforcement comes near to zero, conditioned reflexes will
be progressively eliminated, because they lose their significance.
• The conditioned reflex precedes the unconditioned one, so it has
signalling significance – they prepare the organism to the purposeful
behavior (to avoid the danger, to catch a prey, etc.)
• Thereby,
conditioned reflexes are individual acquired
systemic adaptive reactions of an organism that arise
as a result of temporal-spatial connection in the cortex
hemispheres between the conditioned (sign) irritant
and unconditioned reflex representation
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 Classification of conditioned reflexes
• 1. Conditioned reflexes are named according the name of
unconditioned reflexes on which base they was produced: food,
defense, sexual, orientation
• 2. According the receptors location two large group of
conditioned reflexes are divided:
• Exteroreceptive and interoreceptive.
• Among the exteroreceptive conditioned reflexes according the
modality of conditional stimulus such reflexes are designed:
• Visual, Auditory, Olfactory, Gustatory, Tactile, Thermal.
• Interoreceptive conditioned reflexes are formed when the
irritation of visceral receptors is combined with any unconditioned
reflex.
• 3. According the effector link of reflex arch two groups of
conditioned reflexes are divided:
• 1) autonomic (salivation, respiration, papillary), 2) motor.
• 4. According the method of formation (eliciting, elaboration –
выработка) conditioned reflexes can be divided onto two large
groups:
• 1) classical, 2) instrumental

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 Classification of conditioned reflexes
• 5. According the nature of conditioned stimulus:
• Natural - are based on the natural sign of unconditioned
stimulus (for instance, sight and smell of meat, sight of lemon).
• Artificial - reflexes are formed in response to the stimuli that are
not direct.
• 6. According the temporal peculiarities of conditioned and
unconditioned signals interaction.
• 1) available conditioned reflexes – in the case when
conditioned stimulus and reinforcement are represented at
the same time:
• a) coinciding – reinforcement is added to the signal stimulus
at the same time
• b) postponement – reinforcement is after 30 seconds.
• c) delayed – reinforcement is after 1-3 minutes.
• 2) trace conditioned reflexes are formed when reinforcement
arises after termination of conditional stimulus action.
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 DYNAMIC STEREOTYPE
The chain of stereotypical reactions (may comprise either
conditioned or unconditioned reflexes), which is
developed during individual life in response to constant
sequence of irritants, is called the dynamic stereotype

The mechanism:
1) the sequence of irritants that are
regularly repeated during long time
produces the chain of stereotypical
conditioned reflexes
2) after consolidation of these
reflexes, the entire complex of
reactions (dynamic stereotype) can
be elicited by a single irritant

In natural conditions the dynamic

stereotype results in formation of
habits and other automatic actions.

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 Inhibition of conditioned reflexes
The functioning of conditioned reflexes mechanisms are based on the
two major nervous processes: excitation and inhibition.
So, during the formation and strengthening of conditional reflex the
role of inhibitory process increases progressively

Types of conditioned reflexes inhibition

External Internal
(unconditioned) (conditioned)

-extinctive
Protective -delayed
(outrageous)
-differentiating
-conditioned inhibitor 26
 External inhibition
External unconditioned inhibition is urgent suppression of
conditioned reflex under the action of the outside external
stimulus which elicits orientation reflex or another one.
• External inhibition doesn’t require special formation. It is
realized without participation of inner structure of reflex arch
• External inhibition arises even during the first presentation of the
signal
Orientation reflex:
• Any sudden outside stimulus carries new information;
• This stimulus important for complete estimation of current
conditions
• It forms another dominant region of excitation in the cortex and
initiates orientation reflex (“What happen?”)
• When this orientation reflex arises, it inhibits other competitive
reflexes
Biological significance:
• When external inhibition suppresses current conditioned
activity, it allows to the organism switch its attention over to the
new irritant and estimate its importance or degree of
dangerous 27
 Protective (outrageous) inhibition.
This type of conditioned reflex inhibition arises
• 1) under the action of excessively strong stimulus or
• 2) low functional stage of nervous system when the ordinary
threshold irritants are characterized as excessively strong
• The strongest manifestation of the outrageous inhibition is
stupor (total immobility) under the influence of a very strong
stimulus both physical (bomb explosion) or morale nature
(the sudden news about the sickness or the death of the
immediate relatives)
Mechanism of outrageous inhibition:
• formation of local center of excessive excitation;
• generalization of excitation totally in hemispheres cortex;
• according the low of consecutive induction, generalized
excitation is subsequently replaced by the generalized
inhibition
Biological significance: outrageous inhibition protects the brain
from the excessive expenditure of energetic sources (analog
of safety device) 28
 Internal inhibition
Internal (conditioned) inhibition develops gradually without
the unconditioned reinforcement of conditioned stimulus
• It requires a special conditions for its formation, and arises
step-by-step
• The foundation of internal inhibition is modification of inner
structure of the spatial and temporal connection between the
excited cortical areas that take part in the conditional reflex
realization
Biological significance:
• In changeable conditions internal inhibition allows to
eliminate conditioned reflexes that are not important
more – precise adaptation
• Internal inhibition is the foundation of learning process

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 Extinctive inhibition
If the conditioned irritant is represented without unconditioned
reinforcement, after some isolated applications of conditioned
stimulus the response reaction will decrement
• For some time the animal came
to the forest (or another territory)
and got the food.
• The feeding conditioned reflex
has been formed in response to
the sight of the forest
• Then the food sources run out
here
• The animal can’t find the food
now (i.e. it doesn’t receive the
unconditioned reinforcement)
• The sight of place which was the
positive signal became the
negative one
• So, after some visits of this territory
the extinctive inhibition develops
and the animal will leave this
territory and find another place
with reach food sources.
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 Delayed inhibition
• It is observed when
unconditioned reinforcement
CR comes too late (2-3 min) after
conditioned stimulus
2-3 min presentation
UR • So, the excitation of the cortical
center of conditioned reflex
CR disappears (or becomes
significantly weaker) before the
UR center of unconditioned reflex
will be excited
• Thus the internal temporal and
spatial connections between
these two centers of excitation
get broken.

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 Differentiating inhibition
Differentiating inhibition develops gradually in response to
presentation of several similar factors, if one of them is reinforced,
but the rest are not
• First period – stage of generalization. Conditioned reflexes arise in
response to action of all conditioned stimuli that are represented.
• Second period - specialization. Just one center which is
accompanied by the unconditioned reinforcement keeps
excitation and produce the conditioned reflex
• Differentiating inhibition is the foundation of numerous types of
learning (training) to form the fine skills
Example:
reinforcement • In animal or human conditional
+ CS 1 1 reflex has been formed to the
appointed frequency of sound
‒ CS 2 2 stimulus (for instance, frequency
no reinforcement of metronome sound 45, 50 and
55 per second)
UR • When during some time one
stimulus (50 sounds/sec) is
reinforced with unconditional
stimulus, but other similar stimuli
(45 and 55 sounds/sec) are not,
the conditional reflexes to the
last stimuli reduce. 32
 Conditioned inhibitor
• If the conditioned stimulus in
response to which the
conditioned reflex was formed is Example:
in combination with another • Your cat has a food-driving
stimulus and their combination conditional reflex to the look of
is never reinforced with the food on the table
unconditional stimulus, • But if you see how it steals the
conditional reflex that is food, you will punish the pet
induced by this stimulus will be • In this case you are the
inhibited. conditioned inhibitor for the
animal
• The combination of the food on
the table and your presence is
never reinforced by the food
• After some time this
conditioned reflex will be
inhibited and your cat never
will take the food from the table
in your presence
• Although, this isn’t always true
in your absence!
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TYPES OF HUMAN HNA

• Hippocrates has defied

four types of human
temperament:
• Sanguine (blood)
• Phlegmatic (phlegm)
• Choleric (yellow bile)
• Melancholic (black bile)

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 Pavlov’s conception of human types of
temperament is based on the properties of main
processes in the CNS – excitation and inhibition

1. Intensity of main processes – the strength of

excitation or inhibition in the CNS

2. Balance of main processes – what process is

predominant: excitation or inhibition

3. Lability of main processes – ability to go from

excitation to inhibition and back

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 Types of temperament

Strong Weak

Balanced Unbalanced
Weak type
Mobile Inert

Unrestrained,
Energetic Quite
uncontrollable

SANGUINE PHLEGMATIC CHOLERIC MELANCHOLIC

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 Types of temperament
STRONG & BALANCED
SANGUINE
People with this type of temperament
possess strength and mobility of main
neural processes:
• They readily get over the difficulties
• Have good orientation in new
surrounding
• They are self-possessed

PHLEGMATIC
This type is characterized by strength and
balance, but low lability. This people are
• Persistent, stead-fast toilers
• Well-balanced but slow in making
decisions
• Confined in their habits 37
Types of temperament
CHOLERIC
This type is marked by high intensity but
unbalance of main processes. People
belong to this type are:
• Easily carried away, distracted
• Passionate
• Easily and quickly irritated and
pugnacious

MELANCHOLIC
This type is characterized by weakness of
main processes. This people are
• Weak-willed
• Afraid and escape of difficulties
• Easily subjected to the influence of
others
• Always anxious and gloomy 38
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