CHARACTERISTICS

OF EXCITABLE TISSUES.
TRANSPORT OF
SUBSTANCES.
BIOELECTROGENESIS

Editor: Kuzhel O.P.

 he general properties of
excitable tissues (muscular,
nervous, glandular)
irritability, excitability,
conductivity, lability
 General properties of excitable tissues
1. Irritability property of all tissues to answer on
action of weak irritants by short-term, slowly
developing nonspecific reaction (change metabolism).
2. Excitability property of the high-organized tissues
quickly to react to action of stronger irritants by
specific reaction.
3. Conductivity property of excitable tissues to
conduct excitement in the form of discrete acts
(impulses).
4. Lability (functional mobility) is defined by the
maximum quantity of impulses of excitement which the
tissue in unit of time according to an irritation rhythm,
i.e. without rhythm transformation, is capable to
conduct. It is measured in Hz (impulses/second).
 Irritant change of the external or internal
environment arising quickly enough, acting long
enough and being intensive enough.
Classify on force: subthreshold stimulus, threshold
stimulus, suprathreshold stimulus , maximum stimulus
Classify for nature (physical, chemical, biological etc.).
Modality the qualitative characteristic of an irritant (for
example, a sound, light etc.).
Adequate irritants operate under natural conditions, are
perceived by the corresponding receptors.
Inadequate irritants operate in artificial conditions,
force them should be more than adequate. The most important
an electric current.
 Stimulation threshold(S)
the minimum intensity of
an stimulus causing the
b minimum specific answer.
c

As well as lability, a stimulation

threshold is a measure of
excitability ()
E=1/S
 MODERN IDEAS OF THE STRUCTURE AND FUNCTION
OF BIOLOGICAL MEMBRANES
HYDROPHYLIC GLICEROLS

HYDROPHOBIC FAT ACIDS

The membrane consists from phospholipids and proteins.

Layer of phospholipids double. Their hydrophylic parts
(heads) are directed to a membrane surface, and hydrophobic
parts (the tails stabilizing a membrane) into a membrane.
37
channels
transmembrane ways
on which between
cytosol and structural
intercellular space proteins
(and in the opposite
direction) water, ions
and molecules move enzymes
pumps move receptors
ions against their
concentration and carriers carry out
electrochemical transmembrane moving of
gradients by certain molecules (including,
means of the in a combination with the
energy released at transfer of ions or molecules
hydrolysis of ATP of other type)
39
 TRANSPORT OF
SUBSTANCES THROUGH
UNIPORT
THE MEMBRANE
Transport proteins are
specific: transfer through lipid
bilayer, as a rule, one
substance. SYMPORT
Distinguish: uniport C
unidirectional transport O
of one substance; T
symport unidirectional R
A
transport of two different N
substances; S
antiport (exchanger) P
transport of two different O
R ANTIPORT
substances in opposite T
directions.
41
 PROPERTIES of electroexcitable membrane
CHANNELS:
1) selectivity is the selectively increased permeability of
the ion channel for certain ions and lowered for others.
s defined by the selective filter; 2) dependence from
membrane potential gate of the channel are open or
closed depends on potential on a membrane.
CHANNEL PRONEINS
LIPID BILAYER

ION CHANNEL

THE CHANNEL IS OPEN THE CHANNEL IS CLOSED

 WORK OF GATES OF THE SODIUM CHANNEL
REST DEPOLARIZATION INACTIVATION

m gate
activation

h gate inativation, work more

slowly, are located at an internal
opening of the channel
50
 The doctrine about biopotentials is connected with
Alessandro Volta and Luigi Galvani
opponents in the well-known in the history of a science
dispute on an animal electricity.

Luigi Galvani Alessandro Volta

(1737 - 1798) (1745 - 1827)

52
 MEMBRANE POTENTIAL (MP)
or RESTING POTENTIAL E0
This is a difference of charges between external and internal
surfaces of a membrane. In a nervous cell MP=-70 mV.
The difference size scalar, i.e. can't be neither positive, nor
negative. The minus sign specifies that the microelectrode is on
an internal surface of a membrane which is charged negatively.

mV In drawing it is visible
100
that at the moment of a
membrane puncture the
50 beam on the screen of an
oscillograph deviates
0 down.
microelectrode
-50
cell E0
-100

57

The classical:
1) diffusive-ionic
(Chagovets, 1896)
explains formation of
biopotentials by simple
diffusion of ions;
2) membrane-ionic
(Bernstein, 1902) it is
based on selective
permeability of a
membrane and ionic
asymmetry (a difference of
concentration of ions on the
different parties of a
membrane).
ELETROGENESIS THEORIES
The modern: 1) membrane-ionic
(Hodzhkin, Huxley, Katts, 1949 -
1952, are awarded the Nobel Prize). Is
conventional;
2) phase sorbcionic cytoplasmatic
(Nasonov, Aleksandrov, Troshin, the
middle of the twentieth century) salts
is not in solutions, and are distributed
in phases, i.e. connected with the
various organic substances which
sorbcionic properties change at
excitement.
58
 MEMBRANE-IONIC THEORY
Conditions for biopotentials generation:
1. Selective permeability of a membrane passes
one substances and doesn't pass others. Not to
confuse to a semipermeable membrane which
passes only solvent and doesn't pass the dissolved
substance!

59
 In 1949 Alan Hodzhkin and Bernard Katts calculated a
ratio of permeability (g) of a membrane for various ions in
experiences on huge axon of a squid.
In rest ratio gK+:gNa+:gCl- = 1:0,04:0,45.
It means that in these conditions the membrane well
permeable for ions of potassium and is almost
impermeable for sodium ions. Later and on cells
of mammalians it was shown that at rest permeability
of a cellular membrane for K ions in 20100
times is higher, than for Na+ ions.
At excitement gK+:gNa+:gCl- = 1:20:0,45.
Therefore, at excitement permeability of a membrane for
sodium ions sharply increases.
 Second condition ion asymmetry. The intracellular fluid of living
cells, the cytosol, has a composition very different from that of the
extracellular fluid. For example, the concentrations of potassium
ions are higher inside cells than outside (in 30 - 50 times), whereas
sodium (in 10-15 times), calcium, and chloride (in 30 - 50 times) ions
concentrations are much lower inside cells than outside.

Na+

+ Cl -
 The third condition work of the ion pump Na+, K+-ATF-
ase. The pump forces potassium in a cell, and sodium in
extracellular liquid against electrochemical gradients. The number
of transferable ions (q) isn't equal (qNa+/qK+ = 3/2), i.e. the pump
gives rise to a potential difference. Is a direct electrogene effect of
the pump (let's designate it as Epump). Indirect participation of
the pump in creation of concentration gradients.

ADP+Pi

63
 PROCESSES AT MP FORMATION
2. Active: 1) against a
1. Passive diffusion on gradient; 2) with an expense
concentration and of energy; 3) with carrier
electrochemical participation (active transport
gradients. is always specific, i.e. to each
substance own carrier).
ACTION
POTENTIAL (AP)
This is a fluctuation
of MP at excitement
(in a nervous cell
from 70 mV
to +50 mV).
At excitement permeability of a membrane for sodium ions
sharply increases. At the expense of energy of an irritant
the MP changes. Activation gates open, inactivation start
to be closed. But their kinetics slower, therefore some time
the membrane will be permeable for sodium.

m gate
activation

h gate inactivation
Sodium moves to a cell on
concentration and
electrochemical (+ to -)
gradients and will neutralize a
negative charge of an internal
surface of a membrane. The
potential difference
(polarization) decreases
depolarization. When the
difference of charges will
disappear, sodium continues to
move to a cell on a
concentration gradient and
charges an internal surface of a
membrane positively.
77
 ARE OPEN ARE INACTIVATED

ARE
CLOSED
IN THIS DRAWING THE CONDITION OF SODIUM CHANNELS IN VARIOUS
PHASES OF ACTION POTENTIAL IS REPRESENTED
AFTER PEAK OF POTENTIAL OCCURS MP RESTORATION A REPOLARIZATION

2. Increase in
permeability for
potassium (it starts to
raise together with
permeability for sodium,
since potassium
channels too
dependence from
membrane potential, but
their kinetics slower,
than at sodium
channels).
1. A sodium inactivation sharp
decrease in permeability for
sodium (inactivation gate are 3. Work of Na+,+-pump.
closed).

MECHANISMS OF REPOLARIZATION
 mV
+50

Besides a current of rest and a

current of action exist spike
biopotentials:
1. Local answer.
2. After potentials: negative
(repolarization delay) when
is inhibited a sodium negative after
-50 potential
inactivation, positive
(hyperpolarization) when local
permeability for potassium response

increases. -70

3. Postsynaptic potentials
exciting and inhibiting. positive after
potential
 FEATURES OF LOCAL AND EXTENDING
EXCITEMENT
LOCAL EXTENDING
EXCITEMENT EXCITEMENT

force of stimulus 50 - 75 % from the threshold the threshold

ability to distribution doesn't extend or extends extends
with decrement on 1 - 4 mm

ability to a summation it is capable to a summation it is not capable to a

summation

dependence from force of direct dependence doesn't depend (submits to

stimulant the law "everything or
nothing"
excitability raises Decreases

duration is longer, than action is less long, than local

potential excitement
86
RATIO OF EXCITABILITY mV

PHASES WITH PHASES OF +50

ACTION POTENTIAL.
At the moment of the local answer
spike
excitability raises, at the moment PHASES OF
ACTION
of a rising phase of peak POTENTIAL
decreases (an absolute refractory
period on any irritants the cell -50
negative after
potential
doesn't answer). local
At the moment of a falling phase of answer
peak excitability is restored (a -70

relative refractory period the cell

can answer to superthreshold positive after
stimulus). During negative after potential
EXCITABILITY
potential excitability increases.
During positive after potential
excitability decreases. 88
 THE REFRACTORY PERIOD TIME AFTER
GENERATION OF ACTION POTENTIAL DURING
WHICH EXCITABILITY OF THE MEMBRANE
DECREASES, AND THEN IS GRADUALLY RESTORED
TO INITIAL LEVEL

The relative refractory

The absolute refractory
period an interval
during which the
excitable tissue isn't
capable to generate
action potential,
whatever strong was an
irritant
8
period an interval
during which the tissue
gradually restores a
excitability and can
answer action of
suprathreshold stimulus
by a generaration of
action potential