Chronobiology:
dialectic of the temporal and the biological
Jeremy
Campbell indicates that Einstein had removed time and space from their traditional
metaphysical pedestal of unchanging absoluteness. In other words, the effect
of relativity theory was to physicalize space and time. As a result, time and space became
fluctuating components of the physical universe capable of entering into dynamic
interactions with other facets of that universe.
Just as time
was physicalized through the efforts of Einstein, Campbell contends time has been "
biologized and psychologized" through the work of a variety of recent
experiments and explorations. According to Campbell, just as Einstein seemed to show that
time interacted with the motion of a given system, biologists have been introducing
experimental data indicating biological clocks are affected by the conditions of life
which surround such clocks.
When
Einstein physicalized space and, especially, time, he was culminating, as well as
transforming, a process popularized by Galileo (though this process did not begin with the
latter). Galileo treated time as a continuous and uniform entity which could be
represented by a straight line. Thus, time was construed in a spatialized
manner within a mathematical framework.
As such,
time came to be treated as if it were a fourth spatial direction which is continuous in
the same way that space is supposed to be continuous. In other words, both space and time
were alleged to consist of an infinite number of points, all of which can be mapped on to
the real number line.
Consequently,
the modern conception of time has deviated rather substantially from the idea of time
which had prevailed for nearly 2000 years. In the traditional view, time was considered to
be some sort of absolute master clock which was independent from all of physical/material
reality. Today, time has become just another component of the physical world
which is capable of fluctuating under a variety of conditions.
However, all
of these changes in the way in which time is, and has been, conceived may be more a
reflection of the way time is methodologically engaged than they are a reflection of the
structural character, or actual ontology, of time. In other words, what really may have
changed in the last 2000 years is the way in which time is methodologically engaged.
These
transitions in methodology have led to comparable transformations in the way that time is
conceptualized. None of these changes, however, necessarily has anything to do with giving
insight into the ontology of time.
Zeitgebers:
extrinsic and intrinsic
Organisms
are not only oriented in space, they are also oriented in time. Chronobiology is
the science which studies the role that temporality has in biological functioning. A great deal of relatively recent experimental findings suggests there are innate
mechanisms in a large number of species of organisms which give expression to a variety of
temporal rhythms. These rhythms regulate different facets of biological and behavioral
processes in various species.
For example,
consider animals living in burrows. Such animals have an internal, biological clock which
is entrained by the temporal rhythm of alternating patterns of night and day.
Each day,
the internal, biological clocks of these animals are reset to reflect the changing
relationship of the ratio of daylight hours relative to nighttime hours. When they wake up
in the morning, their internal clocks, not the light of day, has awakened them.?
Franz
Halberg introduced the term circadian rhythm to describe those
instances of temporal entrainment, such as in the case of the burrow animals mentioned
above, that are based on a period lasting roughly one day. The alternating cycle of day
and night acts as a zeitgeber or 'time giver' which an organism uses as
a temporal frame of reference to set its circadian biological clock.
When an
organism is disentrained - that is, when an organism is unable to make contact with the
temporal frame of reference provided by the relevant zeitgeber (in this case, the
alternating cycle of day and night), such a disentrained organism will operate on the
basis of the intrinsic properties of its internal biological clock. This clock, left on
its own without any external standard by which to set itself, will run either somewhat
longer than a 24-hour period, or somewhat shorter than a 24-hour period.
Organisms
entrained by various kinds of temporal rhythms, of which circadian rhythms are but one
example, do more that just reset their internal clocks to synchronize with various rhythms
of the external world. Entrainment means virtually every biological process which goes on
in a given organism will have a determinate phase relationship with events occurring both
in other parts of the body, as well as in various aspects of the external world.
The
phenomenon of diapause is an example of how the behavior of an organism can be
governed by the phase relationships which the biological clock of that organism
establishes with respect to certain features of the external world. Diapause refers to the
period of inactivity or quiescence exhibited by many insects during relatively regularly
occurring periods of detrimental weather conditions, such as drought or winter weather.
However, the
preliminary stages of diapause occur much in advance of the forthcoming, adverse weather
conditions. Insect activities such as the storing of food or the building of shelters are
steps that are preparatory in nature and which take place independently of any specific
stimuli of drought or cold or snow.
The
preparatory activity is an expression of the phase relationships which exist among: (a)
certain biological clocks of the insect; (b) various motor systems in the insect, and (c)
the changing ratio of sunlight to nighttime. As the character of these phase relationships
changes, behavioral patterns emerge which are preparatory to the later set of phase
relationships which constitute diapause proper - that is, the actual period of quiescence.
Therefore,
biological clocks are part of a system which enables an organism to grasp (although not
necessarily on a conscious level or in a self-reflexive manner) the character of a
changing set of phase relationships in the dialectic between organism and environment. In
a sense, there is a process in which certain rules of temporality are internalized.
These rules have the effect of placing constraints on the freedom of an organism to act.
Rules
versus principles
From the
perspective of the present article, the internalization of rules of temporality is not
really an accurate way of describing the situation. More specifically, the organism
consists of a spectrum of ratios of constraints and degrees of freedom. This spectrum
establishes a set of parameters within which, and through which, the organism is capable
of responding or manifesting itself under appropriate circumstances of dialectical
interaction with the environment.
Although
phase information may be exchanged, and although the effect of this exchange of phase
information may bring about a transition in the aspect of the organism's spectrum of
ratios which is being manifested, no rules, temporal or otherwise, are internalized by the
organism. A principle is activated, instead, through the dialectical activity.
The term
"principle" refers to certain kinds of ratios of constraints and degrees of
freedom. Such ratios may be manifested in the form of hermeneutical point-structures,
neighborhoods, or latticeworks (see the Glossary of Terms in the
Overview article within the Education Folder in Food For Thought).
What makes a
given ratio of constraints and degrees of freedom, or set of such ratios, a principle has
to do with the structural character of the phase relationships which exist in the
ratio(s). A principle consists of a set of phase relationships which form an attractor
basin.
The
attractor basin may be either linear or chaotic, depending on the nature of the principle.
However, usually speaking, principles involve chaotic attractors, not linear attractors.
Rules, when
they do arise, tend to be associated with linear attractors. Such attractors are fairly,
narrowly defined and do not permit much, if any, deviation from the scope of the
parameters that describe a rule.
Principles,
on the other hand, provide a basis for a far more sweeping range of possibilities. All
such possibilities are self-similar, rather than self-same.
Consequently,
principles are capable of being receptive to, as well as of responding to, nuances and
variations that fall beyond the largely linear horizons of a rule. Nonetheless, despite
such variability, all these self-similar possibilities fall within the structural
parameters of the chaotic attractor to which they give expression.
The
principle(s) inherent in a given biological clock form an attractor basin which is
sensitive to, and shaped by, certain kinds of phase information being relayed to the
basin(s) as a result of the organism's engagement of, and engagement by, different aspects
of the environment. In other words, the presence of certain kinds of phase relationships
induces shifts or transitions in the way the attractor basin/principle gives expression to
itself. As a result, the principle, in this case a biological clock, is activated.
Subsequent
behavior which is generated in, or which is colored by, such an attractor basin, will
conform to the parameters of constraints and degrees of freedom that have been established
by means of the activated principle/attractor basin. Moreover, since the activated
attractor basin/principle is sensitive to, and shaped by, the changing character of the
phase relationships in the dialectic between organism and the environment, those
behavioral patterns that are influenced by such an attractor will reflect the shifts in
phase relationship information.
In short,
certain aspects of the organism's behavior become entrained by transitions in phase
relationship. Thus, although no rules have been internalized, principles have been set in
motion and behavior has been affected as a result of the dialectical engagement between
organism and environment.
The
question of master biological clocks
In the early
1970s, a certain amount of excitement was generated when a number of biologists believed
they had discovered a master biological clock. Such a clock is supposed to be autonomous
and independent of all external, temporal cues. In addtion, a master biological clock is
theorized to be responsible for generating all the different rhythms of the body.
The would-be
master clock discovered in the 1970s is located in the frontal portion of the
hypothalamus. It consists of several clusters of cell groups which have become linked
during the course of development. The technical term for these coupled cell clusters is suprachiasmatic
nucleus - or, SCN, for short.
Two
properties, in particular, of the SCN seemed to enhance its attractiveness as a candidate
for the master clock. First of all, the coupled nuclei of the SCN display a great deal of
oscillatory activity. Oscillatory behavior is something one would expect to
observe in any candidate for a master clock since the clock is responsible for regulating
a wide variety of rhythmic patterns.
Secondly,
the suprachiasmatic nuclei are connected, via a nerve tract, to the retina in each eye.
One obvious implication of this link is that the SCN would be able to receive important
data concerning temporal rhythms in the external world. Especially important in this
regard would be those rhythms involving the changing pattern of the ratio of daylight to
nighttime as one progressed through the year.
Subsequent
experiments, in which the SCN were removed, indicated the master clock had not been found.
These experiments showed that although the temporal identity of an organism is
significantly altered when the SCN are removed, nevertheless, temporal identity was not
destroyed. In other words, while the SCN seemed to play a fundamental role in synchronizing
various biological rhythms, they were not responsible for generating these
other rhythms. Consequently, there must be other biological sources which are
underwriting temporal identity.
Although the
suprachiasmatic nuclei do not constitute ' the' master biological clock, they are believed
to be the locus within which one of two master clocks can be found. Together, these two
clocks are considered, by many chronobiologists to be responsible for regulating the vast
majority, if not all, of the biological rhythms in the human body. These rhythms range
from: the secretion of growth hormone, to cycles of activity and inactivity, to
establishing the point in the sleep cycle when vivid dreams are most likely to occur, to
the rise and fall of core body temperature, and so on.
The location
of the second master clock has not yet been established. However, this second clock is
thought to be the more stable, as well as the more powerful, of the two clocks.
Nevertheless,
this second, more stable and powerful, master clock is believed not to have any direct
contact with the changing patterns of light to darkness ratios. Therefore, this second
clock may be entrained by the so-called master clock thought to be located in the
suprachiasmatic nuclei, since this latter "master" clock is in contact, via
nerve tracts extending to the retina, with external data concerning the changing ratio of
light to darkness.
There are
some chronobiologists who do not accept the two-master-clock hypothesis. They believe
there may be a number of other "master" clocks in addition to the two already
mentioned.
For example,
there is considerable evidence pointing toward the adrenal gland as the locus for, yet,
another clock of sorts. More specifically, one of the hormones secreted from the outer
cortex of the adrenal glands is cortisol.
Cortisol
plays a fundamental role in the way the body responds to stressful situations.
Fluctuations in the level of cortisol secretion appear to follow cyclical rhythms during
the course of the day.
The
adrenal-clock, however, is not necessarily a master clock. Quite frequently, a given
biological system will have an intrinsic periodicy which characterizes its biological
activity. This innate periodicy is not, in and of itself, a master clock. Such inherently
periodic systems are known as a tau.
The
structural character of a tau gives expression to certain aspects of an underlying genetic
blueprint. Although a tau's general structural character is species specific, the
individual members of a species will display a tau which is similar to, but not precisely
the same as, the average value for the species with respect to that tau.
Human
beings, along with a variety of other species, are capable of being entrained,
simultaneously, to a variety of different biological clocks. On the other hand, human
beings are also capable of having some of their biological rhythms synchronized with
others with whom they live in close contact over a period of time.
Some
hormones play a role in communicating, to various systems in the body, information
concerning the temporal phase of external rhythms. These hormones are referred to as temporally
active hormones.
These sort
of hormones are believed to keep different circadian systems in touch with the
fluctuations occurring in various rhythmic patterns in the external world that are
relevant to the body's circadian rhythms. In human beings, there are a variety of
temporally active hormones providing humans with a number of different sources of temporal
information.
As a result,
such hormones help establish a spectrum of ratios of constraints and degrees of freedom
with respect to the way a human being can engage the environment in a temporal dialectic.
Furthermore, although the general number and structure of biological clocks is pretty much
the same from one human being to the next, there can be a great deal of variance in how
these different clocks are linked together in different individuals. In other words,
different individuals will exhibit different patterns of synchronization with respect to
how the clocks will be linked to one another.
Sometimes
these differences are a result of genetic inheritances. Sometimes the differences in
patterns of synchronization are due to the kind of life the individual leads. Finally,
sometimes a combination of the two foregoing factors will lead to differences in patterns
of synchronization from individual to individual.
A
spectrum of biological rhythms
Modern
high-speed computers have taken on a function, with respect to biological rhythms,
somewhat similar to the role that a prism played with respect to light waves. Just as a
prism is able to show visible light is an aggregate of a number of different wavelengths
of light, so too, modern computers have been able to show there is a spectrum of
biological rhythms underlying an organism's activity.
Through the
application of computer and inferential statistical techniques, approximately seven to
eight basic types of rhythms have been discovered so far. They are: ultradian (less
than 20 hours); circadian (between 20-28 hours); circasemiseptan (31/2
days); circaseptan (7 days, plus or minus 3); circadiseptan (14 days, plus
or minus 3); circavigintan (21 days, plus or minus 3); and, circannual (1
year, plus or minus 2 months). The term infradlan is used to refer to cycles
lasting longer than 24 hours.
Circaseptan
rhythms (which have a period of approximately 7 days) are showing up in a variety of
biological processes. Generally speaking, these rhythms are of low amplitude and,
therefore, are hard to detect amidst the higher amplitude, more prevalent circadian
rhythms. However, although, on an individual basis, the circaseptan rhythms are weaker
than the circadian rhythms, over the course of a week, the aggregate collection of
circaseptan rhythms has a large amplitude.
While
circaseptan and circasemiseptan rhythms do not appear to reflect any external temporal
rhythm, these rhythms are not arbitrary. They have a harmonic relationship with such
external rhythms as the cycle of day and night, as well as the lunar cycle.
Thus, the
rhythms associated with various biological functions (such as growth, maturation, cell
maintenance, reproduction, immune responses, and so on) will be a complex harmonic
function of the way entrainment properties of external rhythms dialectically interact with
the vectoring properties of innate biological currents such as the circaseptan and
circasemiseptan rhythms. However, nobody in the field of chronobiology knows, yet, what
the structural character of this dialectic is or what the harmonic laws are which govern
that dialectic.
One can
differentiate between music and noise by noting how the former consists of a set of sound
waves which have an ordered, structured relationship with one another. In the case of
noise, the aspect of orderly relationship is missing.
In music, a
given complex sound is a function of a set of simple waves which are whole-number
multiples of some fundamental, lowest frequency, wave component inherent in the given
complex sound. This lowest frequency wave component is known as the first harmonic.
Depending on the sort of whole-number multiple a given wave component has relative to the
frequency of the first harmonic, the other wave components of a complex musical sound will
be referred to as harmonics of the second, third, fourth, etc. order.
Some of the
more complex temporal rhythms (e.g., circannual or circavigintan , etc.) may be
whole-number multiples of some of the simpler rhythms such as the ultradian or the
circadian. Thus, the more complex biological rhythms could be seen to be higher order
harmonics of the basic temporal units.
Light,
melatonin and circadian systems
Just as
light plays a fundamental role in Einstein's special theory of relativity, light also
plays a fundamental role in chronobiology. Light is the standard to which the body refers
in order to re-gauge its biological rhythms so they can be synchronized with, among other
things, the primary circadian rhythms generated by the alternating cycle of night and day.
Although
most of the light impinging on the individual's eye is transduced into visual signals, a
certain amount of the light serves as a source of temporal information concerning the
external rhythm of the cycle of day and night. This information is passed on to the
suprachiasmatic nuclei in the hypothalamus. These nuclei are linked with a variety of
other biological clocks and taus. The end result of this dialectic is to permit the
organism to get into an appropriate phase relationship with external rhythms.
The pineal
gland is known as a neuroendocrine transducer. This means it is
capable of converting or translating the action potentials of the nervous system into the
secretion of various kinds of hormones. One of the hormones transduced by the pineal gland
in this fashion is melatonin.
The
suprachiasmatic nuclei is connected to the pineal gland by means of a nerve tract. By
sending certain messages along this nerve tract to the pineal gland, the SCN is able to
control the quantities, and, therefore, activity, of a particular enzyme in the pineal
gland. The enzyme regulated by the SCN plays a role in synthesizing melatonin from a
precursor neurotransmitter, serotonin.
Although the
precise role of melatonin is not presently known, it is deeply implicated in the body's
circadian system which is hooked into external rhythms of night and day. The levels of
melatonin secretion are highest between the hours of 11 at night and 7 in the morning.
Alternatively, the levels of melatonin secretion are lowest during the hours of waking
activity.
Apparently,
light serves as a signal for the suppression of melatonin secretion, whereas nighttime
acts as a stimulus leading to the synthesis of melatonin. The rhythmic rise and fall of
melatonin levels is a waveform which is propagated throughout the body.
This
cyclical waveform plays a role in the synchronization and harmonious interaction of a
variety of biological rhythms. Furthermore, while the amplitude, frequency and phase of
this wave can be affected by altering the timing and/or intensity of the organism's
engagement with light stimuli, each species has its own characteristic way of responding
to such alterations in the character of light stimuli.
Almost all
vertebrates come equipped with a pineal gland. Although the function and the size of the
pineal gland varies from species to species, generally speaking, the more critical the
role(s) which is(are) played by temporal rhythms in a given vertebrate species, the larger
will be the size of that species pineal gland. In addition, in many of, if not most of
these vertebrate species, fluctuations in the level of melatonin synthesis and suppression
in the pineal gland are linked to the way the organism establishes phase relationships
with external cyclical patterns such as day and night, as well as summer and winter.
In the
latter case, the nervous system may have some sort of mechanism for both: (a) keeping
running totals of the ratio of melatonin synthesis to melatonin suppression and, then, (b)
coupling (a) with a process that compares the latter ratio against some innate or learned
(such as through critical periods) standard. This mechanism allows the organism to make
fairly complex preparations for forthcoming seasonal changes.
The
suprachiasmatic nuclei is also linked with the lateral geniculate nucleus. The primary
neurotransmitter propagated along the nerve tract connecting the SCN and the LGN is known
as neuropeptide Y.
In
experiments in which neuropeptide Y has been introduced directly into the SCN, this
neurotransmitter appears to have the effect of resetting the circadian clock of the
suprachiasmatic nuclei in the same manner as if the organism had encountered the darkness
of night. One of the implications of this kind of experiment is as follows. Just as there
are biochemical components which act as carriers of the temporal information of light,
there also may be systems responsible for the generation and regulation of carriers of the
temporal information of darkness.