Biological Temperature Regulation
Biological Temperature Regulation
OCTOBER 3, 2023
SWatikaa Poovannan
1st year BSc Psychology
Temperature Regulation
Homeostasis and Allostasis
- Physiologist Walter B. Cannon (1929) introduced the term homeostasis to
refer to temperature regulation and other biological processes that keep
certain body variables within a fixed range.
o The process is analogous to the thermostat in a house with heating
and cooling systems. Someone sets the minimum and maximum
temperatures on the thermostat. When the temperature in the house
drops below the minimum, the thermostat triggers the furnace to
provide heat. When the temperature rises above the maximum, the
thermostat turns on the air conditioner.
- Similarly, homeostatic processes in animals trigger physiological and
behavioural activities that keep certain variables within a set range. In many
cases, the range is so narrow that we refer to it as a set point, a single value
that the body works to maintain.
- Analogous mechanisms maintain constant blood levels of water, oxygen,
glucose, sodium chloride, protein, fat, and acidity. Processes that reduce
discrepancies from the set point are known as negative feedback.
o Most motivated behaviour can be described as negative feedback:
Something happens to cause a disturbance, and behaviour continues
in varying ways until it relieves the disturbance.
- In a frightening situation, you begin to sweat before you begin the activity that
calls for sweating. To describe these dynamic changes in set points,
researchers use the term allostasis (from the Greek roots meaning “variable”
and “standing”), which means the adaptive way in which the body changes its
set points in response to changes in its life or changes in the environment.
Controlling Body Temperature
- An average young adult spends about 2,600 kilocalories (kcal) daily. About
two-thirds of your energy is used for basal metabolism, the energy you use to
maintain a constant body temperature while at rest.
- Amphibians, reptiles, and most fish are poikilothermic – that is, their body
temperature matches the temperature of their environment.
o They lack physiological mechanisms of temperature regulation such as
shivering and sweating. Nevertheless, they avoid wide swings in body
temperature by choosing their location within the environment.
- Mammals and birds are homeothermic. They use physiological mechanisms
to maintain an almost constant body temperature over a wide range of
environmental temperatures.
o Homeothermy requires energy and therefore fuel, especially for small
animals. An animal generates heat in proportion to its total mass; it
radiates heat in proportion to its surface area.
o Homeothermic animals can use physiological mechanisms to control
body temperature. To cool ourselves when the air is warmer than our
body temperature, we have only one mechanism, which is sweating.
o Species that don’t sweat will instead pant or lick themselves. Sweating,
panting, or licking exposes water, which cools the body as it
evaporates. This mechanism is limited, however: If the air is humid as
well as hot, the moisture will not evaporate. Furthermore, if you lose
more water by sweating than you gain by drinking, your body runs into
other kinds of problems.
- In contrast, several physiological mechanisms increase body heat in a cold
environment.
o One is shivering. Any muscle contractions, such as those of shivering,
generate heat.
o Second, decreased blood flow to the skin prevents the blood from
cooling before it reaches the brain, heart, muscles, and so forth.
o A third mechanism works well for other mammalian species, though not
for humans: They fluff out their fur to increase insulation. (We humans
have an evolutionary relic of that mechanism. We too fluff out our “fur”
by erecting the tiny hairs on our skin — “goosebumps.” Back when our
ancestors had a fuller coat of fur, that mechanism did some good.)
- However, we also use behavioural mechanisms, just as poikilothermic
animals do. In fact, we prefer to rely on behavioural mechanisms when we
can.
o The more we regulate our temperature behaviourally, the less we need
to rely on energetically costly physiological efforts. Finding a cool place
on a hot day is much better than sweating.
o Finding a warm place on a cold day is much better (and smarter) than
standing around shivering.
- Here are a few other behavioural mechanisms of temperature regulation:
o Put on more clothing or take it off. This human strategy accomplishes
what other mammals accomplish by fluffing out or sleeking their fur.
o Become more active to get warmer or less active to avoid overheating.
o To get warm, huddle or cuddle with others. You might be shy about
hugging strangers to keep warm, but many other species are not. For
example, spectacled eiders (in the duck family) spend their winters in
the Arctic Ocean, crowded together, they not only keep one another
warm but also maintain a 20-mile hole in the ice so they can dive for
fish throughout the winter.
The Advantages of Constant High Body Temperature
- We spend about two-thirds of our total energy maintaining body temperature
(basal metabolism).
- A poikilothermic animal, such as a frog, has a lower level of basal metabolism
and consequently needs far less fuel.
- If we didn’t maintain a constant, high body temperature, we could eat much
less and therefore spend less effort finding food.
o Unlike fish a fish which recruits more and more fast-twitch muscle
fibres to remain active at the risk of rapid fatigue when the water gets
colder, birds and mammals, keep their muscles warm at all times,
regardless of air temperature, and therefore stay constantly ready for
vigorous activity.
- A warmer animal has warmer muscles and therefore runs faster and with less
fatigue than a cooler animal.
o However, we have trade-offs. To get even hotter than 37° would
require still more energy. Furthermore, beyond about 40° or 41°C,
proteins begin to break their bonds and lose their useful properties.
Brain Mechanisms
- All the physiological changes that defend body temperature— such as
shivering, sweating, and changes in blood flow to the skin—depend
predominantly on certain areas in and near the hypothalamus, at the base of
the brain.
- The most critical areas for temperature control are the anterior hypothalamus
and the preoptic area, which is just anterior to the anterior hypothalamus. (It is
called preoptic because it is near the optic chiasm, where the optic nerves
cross.)
- Because of the close relationship between the preoptic area and the anterior
hypothalamus, they are often treated as a single area, the preoptic
area/anterior hypothalamus, or POA/AH.
- The POA/AH monitors body temperature partly by monitoring its own
temperature.
o When an experimenter heats the POA/AH, an animal pants or sweats,
even in a cool environment.
o If the same area is cooled, the animal shivers, even in a warm room.
- These responses are not simply reflexive. An animal will also react to a
heated or cooled POA/AH by pressing a lever or doing other work for cold air
or hot air reinforcements.
- Besides monitoring their own temperature, cells of the POA/AH also receive
input from temperature-sensitive receptors in the skin and spinal cord.
- The animal shivers most vigorously when both the POA/AH and the other
receptors are cold; it sweats or pants most vigorously when both are hot.
- Damage to the POA/AH impairs a mammal’s ability to regulate temperature.
o After that kind of damage, mammals are reduced to using just
behavioural mechanisms such as seeking a warmer or colder location.
Fever
- Bacterial and viral infections generally cause fever, an increase in body
temperature.
- The fever is not part of the illness; it is part of the body’s defence against the
illness. When bacteria, viruses, fungi, or other intruders invade the body, it
mobilizes leukocytes (white blood cells) to attack them.
- The leukocytes release small proteins called cytokines that attack the
intruders and also communicate with the brain.
- Some cytokines probably cross the blood-brain barrier; however, the main
route of communication is that cytokines stimulate the vagus nerve, which
sends signals to the hypothalamus to initiate a fever.
- A fever represents an increased set point for the body temperature. Just as
you shiver or sweat when your body temperature goes below or above its
usual 37°C, when you have a fever of, say, 39°C, you shiver or sweat
whenever your temperature deviates from that level.
- Newborn rabbits, whose hypothalamus is immature, do not shiver in response
to infections. If they are given a choice of environments, however, they select
a spot warm enough to raise their body temperature – That is, they develop a
fever by behavioural means.
- Certain types of bacteria grow less vigorously at high temperatures than at
normal mammalian body temperatures, developing a moderate fever
increases an individual’s chance of surviving a bacterial infection.
o However, a fever above about 39°C (103°F) in humans does more
harm than good, and a fever above 41°C (109°F) can be life-
threatening.
Thirst
- Water constitutes about 70% of the mammalian body.
- Because the concentration of chemicals in water determines the rate of all
chemical reactions in the body, the water must be regulated within narrow
limits.
- The body also needs enough fluid in the circulatory system to maintain normal
blood pressure.
- People sometimes survive for weeks without food, but not without water.
Mechanisms of Water Regulation
- Different species have different strategies for maintaining the water they need.
- Your posterior pituitary releases a hormone called vasopressin, which raises
blood pressure by constricting the blood vessels. (The term vasopressin
comes from vascular pressure.)
- The increased pressure helps compensate for the decreased volume.
- Vasopressin is also known as antidiuretic hormone (ADH) because it enables
the kidneys to reabsorb water from urine and therefore make the urine more
concentrated. (Diuresis means “urination.”)
Types of Thirst
Not all thirst is the same. Eating salty foods causes osmotic thirst, and losing fluid,
such as bleeding or sweating, induces hypovolemic thirst. The two kinds of thirst
motivate different kinds of behaviour.
Osmotic Thirst
- The combined concentration of all solutes (molecules in solution) in
mammalian body fluids remains at a nearly constant level of 0.15 M (molar).
- This fixed concentration of solutes can be regarded as a set point, similar to
the set point for temperature. Any deviation activates mechanisms that restore
the concentration of solutes to the set point.
- The solutes inside and outside a cell produce an osmotic pressure, the
tendency of water to flow across a semipermeable membrane from the area
of low solute concentration to the area of higher concentration.
- A semipermeable membrane is one through which water can pass but solutes
cannot.
o The membrane surrounding a cell is almost a semipermeable
membrane because water flows across it freely and various solutes
flow either slowly or not at all between the intracellular fluid inside the
cell and the extracellular fluid outside it.
- Osmotic pressure occurs when solutes are more concentrated on one side of
the membrane than on the other.
o If you eat something salty, sodium ions spread through the blood and
the extracellular fluid but do not cross the membranes into cells.
o The result is a higher concentration of solutes outside the cells than
inside, and the resulting osmotic pressure draws water from the cells
into the extracellular fluid.
o Certain neurons detect their own loss of water and then trigger osmotic
thirst, which helps restore the normal state.
o The kidneys also excrete more concentrated urine to rid the body of
excess sodium and maintain as much water as possible.
- The brain detects osmotic pressure by getting a part of the information from
receptors around the third ventricle.
- Of all brain areas, those around the third ventricle have the leakiest blood-
brain barrier.
o A weak blood-brain barrier would be harmful for most neurons, but it is
helpful for those monitoring the contents of the blood.
- The areas important for detecting osmotic pressure and the salt content of the
blood include the OVLT (organum vasculosum laminae terminalis) and the
subfornical organ (SFO).
- The brain also gets information from receptors in the periphery, including the
stomach, that detect high levels of sodium, enabling the brain to anticipate an
osmotic need before the rest of the body actually experiences it.
- Receptors in the OVLT, the subfornical organ, the stomach, and elsewhere
relay their information to several parts of the hypothalamus, including the
supraoptic nucleus and the paraventricular nucleus (PVN), which control the
rate at which the posterior pituitary releases vasopressin.
- Receptors also, relay information to the lateral preoptic area and surrounding
parts of the hypothalamus, which control drinking.
Hypovolemic Thirst
- Suppose you lose a significant amount of body fluid by bleeding, diarrhoea, or
sweating.
- Although osmotic pressure has not changed anywhere in your body, you need
fluid.
- Your heart has trouble pumping blood up to the head, and nutrients do not
flow as easily as usual into the cells. Your body will react with hormones that
constrict blood vessels.
- Vasopressin is one such hormone; another is angiotensin II.
o When blood volume drops, the kidneys release the enzyme renin,
which splits a portion of angiotensinogen, a large protein in the blood,
to form angiotensin I, which other enzymes convert to angiotensin II.
o Like vasopressin, angiotensin II constricts the blood vessels,
compensating for the drop in blood pressure.
- Angiotensin II also helps trigger thirst in conjunction with receptors that detect
blood pressure in the large veins.
o However, this thirst is different from osmotic thirst because you need to
restore your body fluids, including the salts, and not just water.
o This kind of thirst is known as hypovolemic thirst, meaning thirst based
on low volume. When angiotensin II reaches the brain, it stimulates
neurons in areas adjoining the third ventricle.
o Those neurons send axons to the hypothalamus, where they release
angiotensin II as their neurotransmitter.
o That is, the neurons surrounding the third ventricle both respond to
angiotensin II and release it. This example suggests that the
connection between a neurotransmitter and its function is not at all
arbitrary; the brain uses a chemical that was already performing a
related function elsewhere in the body.
- An animal with osmotic thirst has an increased preference for pure water, but
one with hypovolemic thirst can’t drink much pure water without diluting its
body fluids and changing their osmotic pressure.
o The animal therefore increases its preference for slightly salty water. If
the animal is offered both pure water and salt, it alternates between
them to yield an appropriate mixture.
o If sufficient salt is not readily available, it shows a strong craving for
salty tastes. This preference, known as sodium-specific hunger,
develops automatically as soon as the need exists, even in infant
animals.
o Sodium-specific hunger depends partly on hormones. When the body’s
sodium reserves are low, the adrenal glands produce the hormone
aldosterone, which causes the kidneys, salivary glands, and sweat
glands to retain salt.
o Aldosterone and angiotensin II together change the properties of the
neurons in the nucleus of the tractus solitarius, part of the taste system,
such that they begin reacting to salt in nearly the same way they would
to sugar.
Hunger
Different species use different strategies of eating.
- A snake or crocodile might have a huge meal and then eat nothing more for
months.
- Bears eat as much as they can whenever they can, their main foods—fruits
and nuts—are available in large quantities for only short times.
- Small birds, at the other extreme, eat only what they need at the moment and
store almost no fat at all.
- Humans eat more than we need at the moment, unlike small birds, but we do
not stuff ourselves like bears.
- Choosing which food to eat and how much is an important decision. We have
a wide array of learned and unlearned mechanisms to help in the process.
The Digestive System and Food Selection
- The digestive system’s function is to break food down into smaller molecules
that the cells can use.
- Digestion begins in the mouth, where enzymes in the saliva break down
carbohydrates. Swallowed food travels down the oesophagus to the stomach,
where it mixes with hydrochloric acid and enzymes that digest proteins.
- The stomach stores food for a time, and then a round sphincter muscle opens
at the end of the stomach to release food to the small intestine.
- The small intestine has enzymes that digest proteins, fats, and carbohydrates.
It is also the site for absorbing digested materials into the bloodstream.
- The blood then carries those chemicals to body cells that either use them or
store them for later use.
- The large intestine absorbs water and minerals and lubricates the remaining
materials to pass as faeces.
Enzymes and Consumption of Dairy Products
- Newborn mammals survive at first on mother’s milk. As they grow older, they
stop nursing for several reasons:
o The milk dries up, the mother pushes them away, and they begin to try
other foods.
o Also, most mammals at about the age of weaning lose the intestinal
enzyme lactase, which is necessary for metabolizing lactose, the sugar
in milk. From then on, milk consumption causes stomach cramps and
gas.
- Adult mammals can drink a little milk, as you may have noticed with a pet dog,
but generally not much. The declining level of lactase may be an evolved
mechanism to encourage weaning at the appropriate time.
- Humans are a partial exception to this rule. Many adults have enough lactase
levels to consume milk and other dairy products throughout life.
- Worldwide, however, most adults cannot comfortably tolerate large amounts
of milk products.
- Most human beings, after all, are Asians, and nearly all the people in China
and surrounding countries lack the gene that enables adults to metabolize
lactose.
- They can eat cheese and yoghurt, which are easier to digest than milk, and
small quantities of other dairy products, but they develop cramps or gas pains
if they consume too much.
- Consequently, they generally avoid dairy products.
Other Influences on Food Selection
- For a carnivore (meat eater), selecting a satisfactory diet is relatively simple.
A lion won’t get vitamin deficient unless it eats vitamin-deficient zebras.
- However, herbivores (plant eaters) and omnivores (those that eat both meat
and plants) must distinguish between edible and inedible substances and find
enough vitamins and minerals.
o One way to do so is to learn from the experiences of others. For
example, children acquire their culture’s food preferences, especially
the spices, even if they do not like every food their parents enjoy.
- There is a phenomenon known as conditioned taste aversion.
o It is a robust phenomenon that occurs reliably after just a single pairing
of food with illness, even if the illness came hours after the food. In
fact, you will come to dislike a food that is followed by intestinal
discomfort even if you know that the nausea came from a thrill ride at
the amusement park.
Short- and Long-Term Regulation of Feeding
Oral Factors
- If you could get all the nutrition you needed by swallowing a pill most of us
would not. Never mind how much time we would save; we like to eat. In fact,
many people like to taste and chew even when they are not hungry.
- In one experiment, college students consumed lunch five days a week by
swallowing one end of a rubber tube and then pushing a button to pump a
liquid diet into the stomach (Jordan, 1969; Spiegel, 1973). (They were paid for
participating.)
o After a few days of practice, each person established a consistent
pattern of pumping in a constant volume of the liquid each day and
maintaining a constant body weight. Most found the untasted meals
unsatisfying, however, and reported a desire to taste or chew
something.
- The opposite of ingesting without tasting is tasting without ingesting. In sham-
feeding experiments, everything an animal swallows leaks out of a tube
connected to the oesophagus or stomach.
- Sham-feeding animals eat and swallow almost continually without becoming
satiated.
- In short, taste and other mouth sensations contribute to satiety, but they are
not enough by themselves.
The Stomach and Intestines
- In one experiment, researchers attached an inflatable cuff at the connection
between the stomach and the small intestine.
- When they inflated the cuff, food could not pass from the stomach to the
duodenum.
o They carefully ensured that the cuff was not traumatic to the animal
and did not interfere with feeding.
o The key result was that, with the cuff inflated, an animal ate a normal-
size meal and then stopped, even though the food stayed in the
stomach.
o Evidently, stomach distension produces satiety. The stomach conveys
satiety messages to the brain via the vagus nerve and the splanchnic
nerves.
- The vagus nerve (cranial nerve X) conveys information about the stretching of
the stomach walls, providing a major basis for satiety.
- The splanchnic nerves convey information about the nutrient contents of the
stomach.
- However, the stomach is not the only part of the digestive system important
for satiety.
o Later researchers repeated the experiment with the inflatable cuff and
replicated the result that a rat stopped eating when the stomach filled,
confirming that stomach distension is sufficient for satiety.
o However, when the cuff was open, much food passed to the duodenum
before the end of the meal, so the stomach never reached full
distension.
- The duodenum is the part of the small intestine adjoining the stomach; it is the
first digestive site that absorbs a significant amount of nutrients.
- Insulin levels rise as someone is getting ready for a meal; the insulin lets
some of the blood glucose enter the cells in preparation for the rush of
additional glucose about to enter the blood.
- Insulin increases even more during and after a meal.
- As time passes after a meal, the blood glucose level falls. Insulin levels drop,
glucose enters the cells more slowly, and hunger increases.
- High levels of insulin generally decrease appetite.
- If the insulin level stays constantly high, the body continues rapidly moving
blood glucose into the cells, including the liver cells and fat cells, long after a
meal.
- Consequently, blood glucose drops and hunger increases in spite of the high
insulin level.
- If the insulin level remains constantly low, as in people with diabetes, blood
glucose levels may be three or more times the normal level, but little of it
enters the cells.
- People and animals with diabetes eat more food than normal because their
cells are starving, but they excrete most of their glucose unused, and they
lose weight.
Leptin (Monitoring Fat)
- Leptin is a peptide that is responsible for fat management.
- Leptin signals the brain about the body’s fat reserves— a long-term indicator
of whether to increase or decrease eating.
- A meal can also increase the release of leptin, so the amount of circulating
leptin indicates something about short-term nutrition as well.
- When leptin levels are high, animals act as if they have plenty of nutrition.
o They eat less, become more active, and increase the activity of their
immune systems.
- If you have enough fat supplies, you can devote energy to your immune
system.
o If you have no fat, you are starving and you have to conserve energy
wherever you can.) In adolescence, a certain level of leptin triggers the
onset of puberty.
o If your fat supply is too low to provide for your own needs, you don’t
have enough energy to provide for a baby.
o On the average, thinner people enter puberty later.
Brain Mechanisms
The Arcuate Nucleus and Paraventricular Hypothalamus
- The arcuate nucleus of the hypothalamus has one set of neurons sensitive to
hunger signals and a second set sensitive to satiety signals.
- The hunger-sensitive cells receive input from the taste pathway. (good tasting
food stimulates hunger)
- Another input to the hunger-sensitive cells comes from axons releasing the
neurotransmitter ghrelin.
o The stomach releases ghrelin during a period of food deprivation,
where it triggers stomach contractions.
o Although ghrelin does not cross the blood-brain barrier, certain brain
neurons release it as a transmitter at the same time the stomach
releases it as a hormone.
- Input to the satiety-sensitive cells of the arcuate nucleus includes signals of
both short-term and long-term satiety.
o Distention of the intestines triggers neurons to release the
neurotransmitter CCK, a short-term signal.
o Blood glucose (a short-term signal) increases secretion of the hormone
insulin. Some neurons also release a smaller peptide related to insulin
as a transmitter.
o Body fat (a long-term signal) releases leptin, a signal of body fat
reserves.
o Insulin and leptin combine their effects onto the satiety-sensitive cells.
- Much of the output from the arcuate nucleus goes to the paraventricular
nucleus of the hypothalamus.
o The paraventricular nucleus (PVN) inhibits the lateral hypothalamus, an
area important for eating. So, the paraventricular nucleus is important
for satiety.
- Axons from the satiety-sensitive cells of the arcuate nucleus deliver an
excitatory message to the paraventricular nucleus, releasing the transmitter
a-melanocyte stimulating hormone (aMSH), which is a type of chemical called
a melanocortin.
o People who have deficiencies in their melanocortin receptors fail to
respond to satiety signals and consequently overeat.
- Input from the hunger-sensitive neurons of the arcuate nucleus is inhibitory to
both the paraventricular nucleus and the satiety-sensitive cells of the arcuate
nucleus itself.
o The inhibitory transmitters here are a combination of GABA (the brain’s
main inhibitory transmitters and two others that are used mainly in the
feeding circuit: neuropeptide Y (NPY) and agouti-related peptide
(AgRP).
o Activity of these transmitters blocks the satiety actions of the
paraventricular nucleus, in some cases provoking extreme overeating.
- An additional pathway leads to cells in the lateral hypothalamus that release
orexin, also known as hypocretin.
o Orexin motivates a near-starving animal to search for food, but less
effect on most meals.
o Insulin, diet drugs, and other procedures that affect eating produce
their effects largely by altering input to the melanocortin receptors.
The Lateral Hypothalamus
- Output from the paraventricular nucleus acts on the lateral hypothalamus,
which includes many neuron clusters and passing axons.
o The lateral hypothalamus controls insulin secretion, alters taste
responsiveness and facilitates feeding in other ways.
- The lateral hypothalamus contributes in several ways –
o Axons from the lateral hypothalamus to the NTS (nucleus of the tractus
solitarius), part of the taste pathway, alter the taste sensation and the
salivation response to the tastes. In short, when the lateral
hypothalamus detects hunger, it sends messages to make the food
taste better.
o Axons from the lateral hypothalamus extend into several parts of the
cerebral cortex, facilitating ingestion and swallowing and causing
cortical cells to increase their response to the taste, smell, or sight of
food.
o The lateral hypothalamus increases the pituitary gland’s secretion of
hormones that increase insulin secretion.
o The lateral hypothalamus sends axons to the spinal cord, controlling
autonomic responses such as digestive secretions. An animal with
damage to the lateral hypothalamus has trouble digesting its foods.
Medial Areas of the Hypothalamus
- A large lesion centred on the ventromedial hypothalamus (VMH) leads to
overeating and weight gain
- Some people with a tumour in that area have gained more than 10 kg per
month.
- These symptoms have been known as the ventromedial hypothalamic
syndrome; damage limited to the ventromedial hypothalamus does not
consistently increase eating or body weight.
- To produce a large effect, the lesion must extend outside the ventromedial
nucleus to invade nearby axons, especially the ventral noradrenergic bundle.
- Rats with damage in and around the ventromedial hypothalamus show an
increased appetite compared to undamaged rats of the same weight after
they gain much weight, they become picky eaters.
o If their diet is bitter or not tasty, they eat less than normal. However,
with a sweetened diet, they eat far more than normal.
o One reason is that they have increased stomach motility and
secretions, and their stomachs empty faster than normal. The faster
the stomach empties, the sooner the animal is ready for its next meal.
o Another reason for their frequent meals is that the damage increases
insulin production so much of each meal is stored as fat. The high
insulin levels keep moving blood glucose into storage even when the
blood glucose level is low.