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SYSTEMOGENY
Function: Traditional and
Systemic
Instead of regarding,
e.g.,views
“vision”, “audition”,
“attention”, “motor control” and “motivation” as
functions, a fundamentally different definition of
a “function” was elaborated in functional system
theory. In this theory, function refers to an
achievement of a result through the
realization of a certain functional system.
Such a function cannot be attributed not only to
certain neural networks but even to the brain.
It relates to the whole organism.
Systemogenesis versus Organogenesis
Systemogeney and Ontogeny
During
early
ontogeny,
those
differently
localized
elements
undergo selective and accelerated
maturation that is essential for
achieving the results of the systems,
providing for the survival of an
organism at the early stages of
individual development.
• Systemogeny takes place not
only during the early ontogenetic
period, but also during adult
development
• The formation of a new
behavioural act is always a
formation of a new system
The essence of the basic idea underlying the
absolute majority of studies within the
framework of the ABC paradigm
A sso cia tion ism  B e h aviorism  C on n ectio n ism
A B C le arning (D .C .D e n ne tt)
A n early illustration of a reflex ,
from D escartes' L'hom m e (1664)
Cartesian concept of animal
spirits
"The traces exist because those
pores in the brain that a spirit
passed through before are more
likely to open when it approaches
the again. And then the spirit can
pass through these pores more
easily."
Long-Term Potentiation and Evoked Spike Responses
in the Cingulate Cortex of Freely Mobile Rats
A. G. Gorkin, K. G. Reymann,
and Yu. I. Aleksandrov
Neuroscience and Behavioral Physiology, Vol. 33, No. 8, 2003
Long-term potentiation of EP
in the cingulate cortex after
tetanic stimulation of the
subiculo-cingulate tract. The
ordinate shows the EP rise
rate, % of initial, and the
abscissa shows time, h. The
time during which posttetanic EP was significantly (t
test, p < 0.05) different from
initial is noted by the line and
asterisks.
Re-juvenilization
Nowadays it is commonly accepted that
many regularities of modification of
functional
and
morphologic
characteristics of neurones, as well as
of control of gene expression, serve as
a basis for the formation of adaptive
behaviour in adults, and are
comparable to those found at the early
ontogenetic stages.
Early genesLate
genesMorphological modifications
 Neuronal death that follows seizures, cerebral ischemia, etc is
initiated by overactivation of glutamate receptors, and an increased
intracellular concentration of Ca2+. Increased concentration of Ca2+
causes transcriptional activiation of so-called “cell-death genes”.
Immediate-early genes play a central role in converting
extracellular signals into long-lasting alterations in cellular
phenotype.
 The existing data provide strong evidence for a relationship
between PERSISTENT activation of immediate-early genes and
delayed neuronal death (that is a form of programmed cell death, or
apopotosis). A similar relationship between persistent expression of
c-fos, and cell death in neural and non-neural systems was apparent.
 By contrast, TRANSIENT induction of immediate-early genes
seems to occur in neuronal populations that are destined to survive.
SS. Schreiber & M. Baudry Trends Neurosci., 1995, 18, 446.
It can be suggested that the expression
of c-Fos is a prerequisite that allows
other transcription components carry
out their specific decision for death
or survival. (p.2736)
Lee Y., Park K.H., Baik S.H., Cha Ch.I. Attenuation of c-Fos basal expression in the
cerebral cortex of aged rat. NeuroReport, 1998, 9, 2733-2736
Compensation
NEONEUROGENESIS
?
Primary and/or
Secondary
assortment?
Pathological
state
??
LG
EG
Secondary assortment
or
Systemogenesis
Systemic specialization
Normal state
Learning
Adults Early onotogeny
Maturation
Primary assortment
Systemic pre-specialization
death
?
DG
EG
EG
Neuron
Neuron is always active
Each step in neuronal elimination is
active.
Neurons of different classes express
receptors whose activation can lead
to their death.
C. Raoul, Pettmann B. & Henderson C.E. Active killing of neurons during development and following stress. Current
Opin. In Neurobiol. 2000, 10, 111-117
Altruism and Suicide
Altruism and social cheating in the social amoeba
Dictyostelium discoideum. Nature, 2000, 408, 965
Strassmann J.E., Zhu Y. & Queller D.C. Most of the time, D. discoideum lives as haploid, free-living,
amoeboid cells that divide asexually. When starved, 104-105 of
these cells aggregate into a slug. The anterior 20% of the slug
altruistically differentiates into a non-viable stalk, supporting
the remaining cells, most of which become viable spores.
If aggregating cells come from multiple clones, there should
be selection for clones to exploit other clones by contributing
less than their proportional share to the sterile stalk. Half of
the chimaeric mixtures show a clear cheater and victim. Thus,
unlike the clonal and highly cooperative development of most
multicellular organisms, the development of D. discoideum is
partly competitive, with conflicts of interests among cells.
Four deaths and a funeral: from
caspases to alternative
mechanisms. Nature Reviews,
2001, 2, 1-10
Leist M. & Jaattela M.
Cell suicide in the adult
nervous system has
serious implications
for the whole
organism
Selection instead of Instruction
 Both in biology and in cognitive sciences there is a principled
discontinuation of the search for a meager stock of all-purpose innate
“primitives”, out of which everything else ought to be “constructed”
through a process of “enrichment”, mediated by “association” …
 The present trend is to grant a very rich innate repertoire and then
look for the mechanisms of internal selection.
 The key role of the environment is not that of supplying more
structure, but rather that of steering a selection.
 We should conceive of “learning” as a process of triggering,
filtering, parameter-setting, progressive specialization and selection.
A SELECTIVE THEORY IS BY NOW ALWAYS
THE MOST PLAUSIBLE CANDIDATE FOR
EVERY PROCESS OF ACQUISITION
[Piattelli-Palmarini M. Evolution, selection and cognition: From “learning” to
parameter setting in biology and in the study of language. Cognition, 1989, 31, 1-44]
The system-selective concept of
learning
“Place-cells” in rats
“Face neurons” in monkeys
“Face neurons” in human beings
«Gnostic neurons» by Konorsky
New Neurons
Studies of information storage in model neural networks
have shown that adding new learning sequentially to the
network can result in "catastrophic interference"
[McCloskey M. & Cohen N.J., 1989]. New information
can obscure previously stored information.
If new memories got new neurons with new connections,
catastrophic interference might be avoided.
(W.T.Greenough et al., Nature Neurosci., 1999)
Studies in our laboratory supported the suggestion that the
mechanism of the formation of behaviour (learning) is the
recruitment (specialization) of “new” (previously silent) neurons
into the subserving of specific new behaviours.
Data from other laboratories (Jog et al. 1999, Thompson & Best
1990, Wilson & McNaughton 1993, Swadlow & Hicks1997,
Woodward et al. 1998) confirms the suggestion
that learning involves “new” neurons rather
than "re-learning" (the replacement of preexisting specialization by a new one) of the
“old” cells;
that newly formed unit specialization remains the same during
the whole period of the recording (weeks and even months),
and that there are many silent neurons in different brain areas
that may become active at some time.
Immune Cells Never Forget
Once learned, some abilities, such as swimming or
riding a bike, are never forgotten even after years
without practice. Others, say running a marathon,
need a regular brushing up. Immunologists have
long debated which category our immunological
memory falls into. Once immune cells learn to
recognize a particular antigen, such as a viral
protein, do they need constant reminders to stay on
top of things, or are their memories permanent?
Two reports in this week's issue of Science … now
bolster the notion that immune cells never forget.
… Most experts are convinced that memory T cells
don't need constant stimulation … to stay in shape
Michael Hagmann Science vol. 286, # 5444, 1999: 1266 - 1267
Konorski’s Concept of Gnostic
Areas and Units
 Unitary perceptions are mediated by the discharge of single
neurons, called gnostic units.
 Gnoctic areas are considered as files of gnostic units
representing all the unitary perceptions established in a
given subject.
 Once a potential gnostic unit had been preemted by a
particular stimulus pattern so as to become transformed
into an actual gnostic unit representing that unitary
perception, it become resistant to any new stimulus pattern.
[Konorski J. Integrative activity of the brain. An interdisciplinary approach. Univ.
Chicago Press: Chicago, 1967, 531 p.]
BEHAVIORAL SPECIALIZATIONS OF
NEURONS
Retrosplenial cortex
DISTRIBUTION OF
FOSFOS-POSITIVE NEURONS
Motor cortex
25
*
10%
3
Retrosplenial cortex
*
20
Motor cortex
3
56%
53%
1
37%
2
5%
Fos-positive cells
1
2
(% of total in the areas)
39%
15
10
5
0
- unidentified neurons
- non-specifically task-related neurons
- specifically task-related neurons
O
«C
L»
RO
NT
T
SI
UI
CQ
«A
IO
N»
T
N- I
O UIS
«N CQ
A
IO
N»
«M
E
NT
AI
NC
NA
E»
*p<0.05 versus the motor cortex.
Intensity of c-Fos expression is the highest in the retrosplenial
cortex which has a higher number of specific behaviorally
specialized neurons
RELATIOSHIPS BETWEEN c-FOS
EXPRESSION AND BEHAVIORAL
SPECIALIZATION OF NEURONS
Retrosplenial cortex
100
m
0.23%
bar-pressing
related neurons
23.6%
Motor cortex
100 m
0.04%
3.6%
0.98%
out of Fos-positive
1.11%
NeoNeurogenesis
 Newly generated neurons in the adult are not only affected
by the formation of a hippocamapl-dependent memory,
but also participate in it.
 The new cells are about 1-2 weeks of age when they
become involved in the learned response.
 This ability to undergo rapid structural change may be a
characteristic of immature neurons that makes them
ideally suited for forming associations between stimuli.
 Newly generated neurons may not be used for learning
under more lenient conditions, but become involved as
task demands increase.
[T.J. Shors, G. Miesegaes, A. Beylin, M. Zhao, T. Rydel, and Gould E. Neurogenesis
in the adult is involved in the formation of trace memories. Nature, 2001, 410,
372-375]
NEW
SYSTEMS
Behavioral Act 2
OLD
SYSTEMS
Individual development
Behavioral Act 1
The new system is
added
to
the
existing systems;
newly
formed
systems do not
substitute
for
previously existing
systems, but are
"superimposed"
over them.
What does it mean - "to superimpose, but not to
substitute"?
Many experiments have demonstrated
that a complex instrumental behaviour
is mastered not only through the
realization of new systems, that were
formed during the process of learning
the acts comprising the behaviour, but
also by the simultaneous realization
of older systems, that had been
formed at previous stages of individual
development.
3
2
REAR WALL
3
2
1
FRONT WALL
4
4
1
8
6
7
AP
PP
5
AF LIF FS
8
7
FG
LIF
6
RC
5
5
0
1.0
1.0
.8
.8
.6
.6
.4
.4
.2
.2
Behavioral Act 2
NEW
SYSTEMS
Behavioral Act 1
OLD
SYSTEMS
Individual development
1 sec
1 2 3 4 5 6 7 8 9 10
1 2 3 4 5 6 7 8 9 10
OLD
SYSTEMS
Individual development
Behavioral Act 1
NEW
SYSTEMS
Behavioral Act 2
OLD
SYSTEMS
Individual development
NEW
SYSTEMS
Behavioral Act 1
Behavioral Act 2
500 msec
Example of the activity of the
M-neuron that was
activated during
rightward body
movement. Activations of
this neuron appear during
the turning to the right
when approaching the
pedal on one side of the
cage (FC), side of the cage
(RC) or during this
movement in defensive
behavior (DB).
NEW
SYSTEMS
Behavioral Act 2
OLD
SYSTEMS
Individual development
Behavioral Act 1
The
realization
of
behaviour
is
the
realization
of
the
history of behavioural
development, that is, of
many systems, each
fixing a certain stage of
development of the given
behaviour.
Behavior is a simultaneous realization of a set of system
ranging from the most archaic to the newest ones
Hippocampus
Position
EMG
Actogr. rear
NEW
SYSTEMS
Behavioral Act 1Behavioral Act 2
Neuron
Neuron
Actogr. front
Actogr. rear
Neuron
Actogr. front
Actogr. rear
OLD
SYSTEMS
Motor cortex
from a hand
from the floor
Trigeminal mesencephalic nucleus
Neuron
Jaw mov.
EMG
500 msec
Individual development
Actogr. front
Memory
Environmental
afferentation
Parameters
of the result
Acceptor
of action’s
result
Decision
making
Triggering
stimulus
Environmental
afferentation
Program
of the action
Result
of the action
Action
Motivation
Afferent
synthesis
r
1
r
R
n
2
Act n
r
1
T
r
2
Act n+1
r
R
n + 1
3
r
1
T
r
2
Act n+2
r
3
RE -CONSOLIDATION the modifications of preexisting memory
Definitive behavior
mismatch
MEMORY
Re-consolidation
in consequence of
memory reactivation
LEARNING
Behavioral Act 2
• SPECIALIZATION - the
recruitment of new neurones
into the subserving of
specific new behaviours
• ACCOMMODATIVE RE CONSOLIDATION - the
modifications of pre-existing
behaviours neurones
NEW
SYSTEMS
Behavioral Act 1
OLD
SYSTEMS
Individual development
TWO CLOSELY CONNECTED PROCESSES:
Memory “projection” to cerebral
structures
Patterns of neurons’ behavioral specialization in different brain
structures
%
100
NEW
60
40
20
0
LimbC Hpc VisC S-sC Hpt MotC OptT
Act 1
Act 2
OLD
SYSTEMS
UNIDENTIFIED
NEW
SYSTEMS
OLD
Individual development
80
Our recordings of unit
activity in many cerebral
structures
during
instrumental
behavior
have demonstrated that
neurons
with
new
specializations
are
abundant in cerebral
cortex,
whereas
phylogenetically archaic
and peripheral structures
contain very few of
them.
E-
E+
28%
Acute
ethanol
11%
Chronic ethanol
HEALTHY
34%
17%
L>M
22%
ALCOHOLIC
34%
M>L
M>L
Acute
ethanol
30%
26%
L~M
The specificity of memory
“projection” to cerebral
structures is determined
by the characteristics of
neurons
composing
these structures. These
characteristics determine
the
involvement
of
neurons of the given
structure
into
the
formation of the certain
behavior.
Brain Research 876 (2000) 154–165
Effects of ethanol on hippocampal place-cell and interneuron activity
* Aaron M. White , Phillip J. Best
Definitive behavior
Mismatch
MEMORY
Re-consolidation
in consequence of
memory reactivation
At the beginning of learning
Pre-exiting
memory
1-EGE
Mismatch
2-Tets
Mismatch
Pre-exiting
memory
3-Success
Mismatch
Pre-exiting
memory
4-«Altruistic suicide»
Pre-exiting memory
5-Consolidation of a new memory
Pre-exiting memory
6-Modification of pre-exiting memory
Pre-exiting memory
New
memory
Accommodative
re-consolidation
7-Dynamics of memory
Pre-exiting memory
Re-consolidation
New
in consequence of
memory
memory reactivation
COGNITIVE SOLIPSISM
 Cognitive science still proceeds as if culture did not matter.
 The central assumption of cognitive solipsism is that the mind may be,
indeed must be, conceptualized as a system that is contained entirely inside a
box. In the case of vertebrates like ourselves, that box happens to be the
brain.
 The strong form of this doctrine holds that [mind] basic structure is a
biological given, structured according to a set of innate neuropsychological
universals. Within this framework, culture is necessarily assigned a
secondary role.
 Lev Vygotsky … was one of the first to recognize the
symbiosis of the developing mind with culture…
[Merlin Donald The central role of culture in cognitive evolution: a reflection on the myth of the “isolated
mind”. In: Culture, thought, and development. L.P.Nucci, G. Saxe, E. Turiel (Eds.) Lawrence Erlbaum
Associates, Inc., 2000, pg. 19-38]
THE UNIT OF CULTURE
We recommended that the unit of culture –
now called meme – be the same as the node
of semantic memory and its correlates in
brain activity
 “Nodes”, or reference points, in semantic
memory … can be associated with neural
activity in the brain
 Nodes are almost always linked to other
nodes, so that to recall one node is to
summon others
 The linkage of nodes is assembled as a
hierarchy to organize information with
more and more meaning
[Edward O. Wilson. Consilience. The unity of knowledge. A.A. Knoff. New
York. 1998, pp. 135,136]
Human specializations and brain
activity
Bird and car experts were tested with
functional magnetic resonance imaging
during tasks with … cars and birds. The
right fusiform girus and occipital lobe
showed significant expertise effect.
[I. Gauthier et al. Expertise for cars and birds recruits brain areas involved in face
recognition. Nature Neurosci. 2000 3 (2), 191 –197]
THE IMPLICATIONS OF NICHE CONSTRUCTION FOR EVOLUTION
Standard Evolutionary Theory
Extended Evolutionary Theory
Laland et al. 1996, 1999, 2001; Odling-Smee et al. 2003
THE IMPLICATIONS OF NICHE CONSTRUCTION FOR HUMAN SCIENCES
Cultural niche construction can influence not only
subsequent cultural changes,
but also some human genetic changes
Gene-culture coevolution
he cultural trait
of dairy
farming is
thought to
have evolved
in association
with the gene
for lactose
absorption