A-Level Biology: Control Systems
A-Level Biology: Control Systems
Contents
Nervous transmission ............................................................ p4
Synapses .................................................................................... p10
Effects of drugs on the nervous system ............................ p13
The Organisation of the Nervous System ........................ p14
The Brain .................................................................................. p18
The Eye ..................................................................................... p21
The Endocrine system ........................................................... p26
Homeostasis............................................................................. p28
Temperature Homeostasis ................................................... p30
Control of Heart Rate ........................................................... p34
Excretion and the Kidney ..................................................... p37
Water Homeostasis ............................................................... p42
Plant Responses ...................................................................... p46
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Stimuli are changes in the external or internal environment, such as light waves,
pressure or blood sugar. Humans can detect at least nine external stimuli and dozens
Stimulus of internal stimuli, so the commonly-held believe that humans have just five senses is
obviously very wide of the mark!
Receptor cells detect stimuli. Receptor cells are often part of sense organs, such as the
ear, eye or skin. Receptor cells all have special receptor proteins on their cell
Receptor
membranes that actually do the sensing, so “receptor” can confusingly mean a protein,
a cell or a group of cells.
The coordinator is the name given to the network of interneurones connecting the
sensory and motor systems. In humans the coordinator is the central nervous system
Coordinator
(CNS) consisting of the brain and spinal cord (p2). Its job is to receive impulses from
sensory neurones and transmit impulses to motor neurones.
Effectors are the cells that effect a response. In humans there are just two kinds:
muscles and glands. Muscles include skeletal muscles, smooth muscles and cardiac
muscle, and they cause all movements in humans, such as walking, talking, breathing,
Effector
swallowing, peristalsis, vasodilation and giving birth. Glands can be exocrine – secreting
liquids to the outside (such as tears, sweat, mucus, enzymes or milk); or endocrine –
secreting hormones into the bloodstream.
Responses aid survival. They include movement of all kinds, secretions from glands and
Response
all behaviours such as stalking prey, communicating and reproducing.
We’re going to be looking at each of these stages in turn, but first we’ll look at the cells that comprise the
nervous system.
Nerve Cells
The nervous system composed of nerve cells, or nerve fibres or neurones. A
neurone has a cell body with extensions leading off it. Several dendrites (or
one dendron) carry nerve impulses towards the cell body, while a single long
axon carries the nerve impulse away from the cell body. Axons and dendrons
are only 10µm in diameter but can be up to 4m in length in a large animal (a
piece of spaghetti the same shape would be 400m long)! A nerve is a discrete
bundle of several thousand neurone axons and dendrons.
Nerve impulses are passed from the axon of one neurone to the dendrites or
cell body of another at a synapse. The dendrites provide a large surface area
for connecting with other neurones.
Most neurones also have many companion cells called Schwann cells, which
are wrapped around the axon many times in a spiral to form a thick lipid layer
called the myelin sheath. The myelin sheath provides physical protection and
electrical insulation for the axon. There are gaps in the sheath, called nodes of
Ranvier, which we’ll examine later.
The combination of the Na+ K+ ATPase pump and the leak channels cause a stable imbalance of Na+ and
K+ ions across the membrane. This imbalance causes a potential difference across all animal cell membranes,
called the membrane potential. The membrane potential is always negative inside the cell, and varies in size
from –20 to –200mV in different cells and species. The Na+ K+ ATPase is thought to have evolved as an
osmoregulator to keep the internal water potential high and so stop water entering animal cells and
bursting them. Plant cells don’t need this pump as they have strong cells walls to prevent bursting (which is
why plants never evolved a nervous system).
The nature of the nerve impulse was discovered by Hodgkin, Huxley and Katz in Plymouth in the 1940s, for
which work they received a Nobel Prize in 1963. They used squid giant neurones, whose axons are almost
1 mm in diameter (compared to 10 µm normally), big enough to insert wire electrodes so that they could
measure the potential difference across the cell membrane. In a typical experiment they would apply an
electrical pulse at one end of an axon and measure the voltage changes at the other end, using an
oscilloscope:
The normal membrane potential of these nerve cells is –70mV (inside the axon), and since this potential
can change in nerve cells it is called the resting potential. When a stimulating pulse was applied a brief
reversal of the membrane potential, lasting about a millisecond, was recorded. This brief reversal of the
membrane potential is actually the nerve impulse, and is also called the action potential:
Since both channels are voltage-gated, they are triggered to open by changes in the membrane potential
itself. The sodium channel opens at–30mV and the potassium channel opens at 0V. The Na+ K+ ATPase
pump runs continuously, restoring the resting concentrations of sodium and potassium ions.
The ion channels have two other features that help the nerve impulse work effectively:
After an ion channel has opened, it needs a “rest period” before it can open again. This is called the
refractory period, and lasts about 2ms. This means that, although the action potential affects all other
ion channels nearby, the upstream ion channels cannot open again since they are in their refractory
period, so only the downstream channels open, causing the action potential to move one way along the
axon.
The ion channels are either open or closed; there is no half-way position. This means that the action
potential always reaches +40mV as it moves along an axon, and it is never attenuated (reduced) by long
axons. In other word the action potential is all-or-nothing.
Synapses
The junction between two neurones is called a synapse. An action potential cannot cross the gap between
the neurones (called the synaptic cleft), and instead the nerve impulse is carried by chemicals called
neurotransmitters. These chemicals are made by the cell that is sending the impulse (the pre-synaptic
neurone) and stored in synaptic vesicles at the end of the axon. The cell that is receiving the nerve impulse
(the post-synaptic neurone) has chemical-gated ion channels in its membrane, called neuroreceptors. These
have specific binding sites for the neurotransmitters.
1. At the end of the pre-synaptic neurone there are voltage-gated calcium channels. When an action
potential reaches the synapse these channels open, causing calcium ions to diffuse into the cell down
their concentration gradient.
2. These calcium ions cause the synaptic vesicles to fuse with the cell membrane, releasing their contents
(the neurotransmitter chemicals) by exocytosis.
3. The neurotransmitters diffuse across the synaptic cleft.
4. The neurotransmitter binds to the neuroreceptors in the post-synaptic membrane, causing the ion
channels to open. In the example shown these are sodium channels, so sodium ions diffuse in down
their gradient.
5. This causes a local depolarisation of the post-synaptic cell membrane, called the post-synaptic potential
(PSP), which may initiate an action potential.
6. The neurotransmitter must be removed from the synaptic cleft to stop the synapse being permanently
switched on. This can be achieved in two ways:
by breaking down the neurotransmitter using a specific enzyme in the post-synaptic cell membrane
(e.g. the enzyme acetylcholinesterase breaks down the neurotransmitter acetylcholine). The
breakdown products diffuse back across the cleft and are absorbed by the pre-synaptic neurone by
endocytosis and used to re-synthesise more neurotransmitter, using energy from the mitochondria.
by absorbing the intact neurotransmitter using an active transport protein in the pre-synaptic
neurone membrane.
These synapses have neuroreceptors that are chloride (Cl- ) channels. When the channels open, negative
Cl- ions diffuse in causing a local hyperpolarisation called an inhibitory postsynaptic potential (IPSP) and
making an action potential less likely. So with these synapses an impulse in one neurone can inhibit an
impulse in the next. Typical neurotransmitters in these synapses are glycine or GABA.
3. Non-channel Synapses.
These synapses have neuroreceptors that are not channels at all, but instead are membrane-bound
enzymes. When activated by the neurotransmitter, they catalyse the production of a “messenger chemical”
(e.g. Ca2+ ) inside the cell, which in turn can affect many aspects of the cell’s metabolism. In particular they
can alter the number and sensitivity of the ion channel receptors in the same cell. These synapses are
involved in slow and long-lasting responses like learning and memory. Typical neurotransmitters are
adrenaline, noradrenaline (NB adrenaline is called epinephrine in America), dopamine, serotonin,
endorphin, angiotensin, and acetylcholine.
4. Neuromuscular Junctions.
These are the synapses formed between effector neurones and muscle cells. They always use the
neurotransmitter acetylcholine, and are always excitatory. Effector neurones also form specialised synapses
with secretory cells.
5. Electrical Synapses.
In these synapses the membranes of the two cells actually touch, and they share proteins. This allows the
action potential to pass directly from one membrane to the next without using a neurotransmitter. They
are very fast, but are quite rare, found only in the heart and the eye.
Synaptic Integration
One neurone can have hundreds (or even thousands) of synapses on its cell body and dendrites. This
arrangement is called axon convergence (because many axons converge on one cell body), and it means the
post-synaptic neurone has many inputs but only one output, through its axon. Each of the many synapses
will produce its own local voltage change, called a postsynaptic potential (PSP), but some of these potentials
will be excitatory (EPSP) and some will be inhibitory (IPSP). The excitatory and inhibitory PSPs from all that
cell’s synapses sum together to form a grand postsynaptic potential (GPSP) in the neurone’s axon. Only if
this GPSP is above a threshold potential will an action potential be initiated in the axon. This process is
called synaptic integration or summation.
Spatial summation is the summing of PSPs from different synapses over the cell body and dendrite tree
Temporal summation is the summing of a sequence of PSPs at one synapse over a brief period of time.
Summation is the basis of the processing power in the nervous system. Neurones (especially
interneurones) are a bit like logic gates in a computer, where the output depends on the state of one or
more inputs. By connecting enough logic gates together you can make a computer, and by connecting
enough neurones together to can make a nervous system, including a human brain.
The beauty of synapses, from a pharmacological point of view, is that synapses in different parts of the
nervous system use different neurotransmitters and neuroreceptors, so drugs can be designed to target
specific synapses and so affect specific aspects of the nervous system. There are three examples of drug
action that you need to understand:
1. Nicotine has a similar structure to acetylcholine, so binds to cholinergic neuroreceptors in the CNS
and switches them on, just like acetylcholine. Nicotine is therefore an agonist of these synapses, but in
an uncontrolled way. The affected neuroreceptors have complex functions involving the release of
other neurotransmitters in the CNS. So by strongly switching on these channels, nicotine causes the
release of more neurotransmitters, including:
adrenaline, which triggers the fight or flight response of the sympathetic nervous system (see p14),
including an increase in heart rate and blood pressure
dopamine, which stimulates the reward system in the brain, leading to feelings of pleasure.
endorphins, which also cause feelings of well-being, but lead to withdrawal symptoms and so cause
addiction.
2. The cobra venom -neurotoxin binds tightly and irreversibly to the acetylcholine receptors in
neuromuscular junctions, stopping the receptors from opening and so stopping skeletal muscles from
contracting. The victim is thus paralysed and is killed if the toxin reaches the breathing muscles.
3. Lidocaine blocks voltage-gated sodium channels, so stopping all action potentials. In low doses
lidocaine has only a local effect at the site of administration, as it is quickly broken down. So lidocaine
acts as a local anaesthetic by inhibiting sensory neurones and as a muscle relaxant by inhibiting motor
neurones. It is commonly used as a quick-acting, short-lasting local anaesthetic, such as in dental
surgery. In high doses lidocaine has more widespread effects and can be fatal.
The spinal cord is enclosed and protected by the 33 bony vertebrae of the spine. The gaps between the
vertebrae allow the spine to flex and also allow branches of the spinal cord to extend out to the periphery
of the body. There are two branches, or roots, on each side of the spinal cord: the dorsal (back) root and
the ventral (front) root. Sensory neurones always enter the spinal cord through the dorsal root, which
contains a swelling, called the dorsal root ganglion, to hold all the sensory neurone cell bodies. Effector
neurones always leave the spinal cord through the ventral root. Their cell bodies are at the end of the cell
in the grey matter of the spinal cord.
Reflexes
The spinal cord transmits nerve impulses between the brain and the periphery through the white matter,
but it also interconnects neurones through the grey matter and so performs simple processing, called
reflexes. A reflex is a specific response to a specific stimulus, which protects the body from damage.
Reflexes are very fast (since they involve few neurones, so few synapses); involuntary (the brain is not
involved); invariable and innate. The path taken by the nerve impulse is called the reflex arc, and involves
only three neurones. Examples of reflexes include the knee-jerk reflex (which protects the knee joint from
damage) and the pupil reflex (which protects the retina from damage by intense light).
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The Brain
The human brain is the site of the major coordination in the nervous system. It contains around 10 11
interneurones, each making thousands of connections to other neurones, so the number of pathways
through the brain is vast. Different regions of the brain can be identified by their appearance, and
techniques such as magnetic resonance imaging (MRI) can be used to study the functions of each region. It
turns out that the regions of the brain have specific different roles.
All the regions except the cerebrum are involved in involuntary functions, and are connected to the
autonomic nervous system. The four regions that you need to know about are:
The Hypothalamus
The hypothalamus controls temperature and water homeostasis (p42). It also controls the release of
hormones by the pituitary gland, including the sex hormones LH and FSH, the kidney hormone ADH, and
human growth hormone HGH.
The Cerebellum
The cerebellum coordinates muscle movement and so controls balance, posture and locomotion (standing,
walking, running, jumping, etc.). The motor areas of the cerebral cortex initiate a particular movement
(such as going for a walk or drawing a picture) but the motor impulses are passed through the cerebellum,
which organises exactly which muscle groups to contract and relax. The cerebellum receives sensory input
from the vestibular system in the inner ear (which senses balance) and from the proprioceptors in the
muscle and joints (which sense body position) and uses this information to coordinate muscle contraction
all over the body to maintain balance and smooth movement. Damage to the cerebellum results in jerky,
uncoordinated movement.
The various functions of the cortex are localised into discrete areas. Some of these areas are shown on this
map of the surface of the left cerebral cortex (you don’t have to learn these).
Receptors
Receptor cells detect stimuli. Humans have over 20 different kinds of receptors. They can be classified as
In some receptors the receptor cell is the sensory neurone itself, while in others, there is a separate
receptor cell that synapses with a sensory neurone. Receptor cells are often part of sense organs, such as
the ear, eye or skin. Receptor cells all have special receptor proteins on their cell membranes that actually
do the sensing, so “receptor” can confusingly mean a protein, a cell or a group of cells. We’ll look at
pressure receptors and light receptors in more detail.
The Eye
The eye is a complex sense organ, not just detecting light, but regulating its intensity and focussing it to
form sharp images. The structure of the eye is shown here.
The actual detection of light is carried out by photoreceptor cells in the retina. The structure of the retina
is shown in more detail in this diagram:
There are two kinds of photoreceptor cells in human eyes: rods and cones, and we shall look at the
difference between these shortly. These rods and cones form synapses with special interneurones called
bipolar neurones, which in turn synapse with sensory neurones called ganglion cells. The axons of these
ganglion cells cover the inner surface of the retina and eventually form the optic nerve (containing about a
million axons) that leads to the brain. Each cone cell is connected to one bipolar neurone, while rod cells
are connected in groups of up to 100 to a single bipolar neurone. This linking together is called retinal
convergence.
A surprising feature of the retina is that it is back-to-front (inverted). The photoreceptor cells are at the
back of the retina, and the light has to pass through several layers of neurones to reach them. This is due
to the evolutionary history of the eye, but in fact doesn’t matter very much as the neurones are small and
transparent. However, it does mean that the sensory neurones must pass through the retina where
converge at the optic nerve, causing the blind spot.
Visual Transduction
Visual transduction is the process by which light initiates a nerve impulse in the retina. The photoreceptor
cells contain hundreds of membrane disks, which each hold thousands of molecules of rhodopsin, the
photoreceptor protein. Rhodopsin is a trans-membrane protein, made up of the polypeptide opsin
surrounding a small molecule derived from vitamin A called retinal.
Retinal is light-sensitive and it can exists in two isomeric forms: a cis form and a trans form. In the dark
retinal is in the cis form, but when it absorbs a photon of light it quickly switches to the trans form, in a
process called bleaching. Rhodopsin with trans retinal changes shape to an unstable form that dissociates
into opsin and free retinal. The opsin initiates a cascade of chemical reactions in the rod cell that eventually
cause an action potential to the brain.
The reverse reaction (trans to cis retinal) requires ATP and several enzyme reactions and is very slow,
taking a few minutes. This explains why you are initially blind when you walk from sunlight to a dark room:
in the light almost all your retinal was in the trans form, and it takes some time to convert enough into cis
retinal to turn off the nerve impulse.
How does the bleaching of the rhodopsin send a nerve impulse to the brain? The details of the process are
complicated and unexpected. Like all neurones, rod cell membranes contain the Na+ K+ ATPase pump,
which generates a sodium gradient. They also have a special sodium channel that is gated indirectly by
rhodopsin.
In the dark the sodium channels are open, so sodium ions diffuse in, so the rod cell is depolarised and
releases neurotransmitter at its synapse. However the synapse with the bipolar cell is an inhibitory
synapse, so the neurotransmitter stops the bipolar cell generating its own nerve impulse, so there are
no impulses to the brain.
In the light rhodopsin with trans retinal initiates a cascade of chemical reactions in the rod cell that close
the sodium channels. Sodium ions stop diffusing in, causing a hyperpolarisation. If enough sodium
channels close the hyperpolarisation reaches a threshold and the synapse is switched off. The bipolar
cell now generates and action potential, which is transmitted to the sensory neurone and so to the
brain.
Fortunately you don’t have to remember these details, but you should be able to understand them.
The rods and cones serve two different functions as shown in this table:
Rods Cones
Shape Outer segment is rod shaped Outer segment is cone shaped
Visual Rhodopsin Photopsin (aka iodopsin)
pigment
Density and 109 cells per eye, distributed throughout the 106 cells per eye, found mainly in the fovea,
distribution retina, so used for peripheral vision. so can only detect images in centre of
retina.
Colour Only 1 type, so only monochromatic vision. 3 types (red green and blue), so are
responsible for colour vision.
Connections Many rods connected to one bipolar cell, Each cone connected to one bipolar cell, so
giving retinal convergence. no convergence.
Sensitivity High sensitivity due to high concentration of Low sensitivity due to lower concentration
(ability to rhodopsin, and to retinal convergence – of rhodopsin and to no convergence – one
detect low one photon per rod will sum to cause an photon per cone not enough to cause an
light intensity) action potential. Used for night vision. action potential. Need bright light, so work
best in the day.
Acuity Poor acuity due to low density in periphery Good acuity due high density in fovea and
(ability to of retina, and retinal convergence (i.e. rods 1:1 connections with interneurones (i.e.
resolve fine are not good at resolving fine detail). cones are used for resolving fine detail such
detail) as reading).
Although there are far more rods than cones, we use cones most of the time because they have better
acuity and can resolve colours. Since the cones are almost all found in the fovea, a 1mm² region of the
retina directly opposite the lens, we constantly move our eyes so that images are always focused on the
fovea. You can only read one word of a book at a time, but your eyes move so quickly that it appears that
you can see much more. The more densely-packed the cone cells, the better the visual acuity. In the fovea
of human eyes there are 160 000 cones per mm2, while hawks have 1 million cones per mm2, so they really
do have far better acuity.
Colour Vision
There are three different kinds of cone cell, each with a slightly different form of opsin, but the same
retinal. These three forms of photopsin are sensitive to different parts of the spectrum, so there are red-
sensitive cones (max absorption at 564nm), green-sensitive cones (max 533nm) and blue-sensitive cones
(max 437nm). By contrast, rods have maximum absorbance at 498nm.
Coloured light stimulates these three cells differently, so by comparing the nerve impulses from the three
kinds of cone, the brain can detect any colour. For example:
Colour Wavelength Red cone absorption Green cone absorption Blue cone absorption
of light (nm) (as % of max) (as % of max) (as % of max)
Cyan 480 12 45 51
Yellow 570 98 62 2
Red 650 10 2 0
So each colour of light has a unique pattern of nerve impulses from the three kinds of cone. The brain uses
this “in reverse” to determine the colour from the pattern of nerve impulses. This is called the
trichromatic theory of colour vision. Colour vision is more complicated than this as most colours are
composed of many different wavelengths and our ability to detect colours also depends on lighting
conditions and other features of the image, and colour vision is a complex interaction of photoreceptors
and neural processing in the retina and in cerebral cortex of the brain.
The opsin proteins in red, green and blue cones are made by three different genes. The blue gene is on
chromosome 7, but the green and red genes are on the X chromosome. This means that males have only
one copy of these two genes (i.e. they’re haploid for these genes). About 8% of males have a defect in one
or other of these genes, leading to red-green colour blindness. Other forms of colour blindness are also
possible, but are much rarer.
Hormones are secreted by glands into the blood stream. There are two kinds of glands:
Exocrine glands secrete solutions to the outside, or to body cavities, usually through ducts (tubes). e.g.
sweat glands, tear glands, mammary glands, digestive glands.
Endocrine glands do not have ducts but secrete chemicals directly into the tissue fluid, whence they
diffuse into the blood stream. e.g. thyroid gland, pituitary gland, adrenal gland. The hormone-secreting
glands are all endocrine glands.
This table shows some of the main endocrine glands and their hormones. The hormones marked with a *
are ones that we shall look at in detail later.
Once a hormone has been secreted by its gland, it diffuses into the blood stream and is carried all round
the body to all organs. However, it only affects certain target organs, which can respond to it. These target
organs have specific receptor molecules in their cells to which the hormone binds. These receptor
molecules are proteins, and they form specific hormone-receptor complexes, very much like enzyme-
substrate complexes. Cells without the specific receptor will just ignore a hormone. The hormone-
receptor complex can affect almost any aspect of a cell’s function, including metabolism, transport, protein
synthesis, cell division or cell death.
There are two different ways in which a hormone can affect cell function:
Most hormones do not enter the cell, but bind The steroid hormones are lipid-soluble so can
to a receptor protein on the cell membrane, to form easily pass through cell membranes by lipid diffusion.
a hormone-receptor complex. This complex then They bind to receptor proteins to form a hormone-
activates an enzyme in the membrane. For example receptor complex in the cytoplasm, which enters
adrenaline activated the enzyme adenylate cyclase, the nucleus and stimulates protein synthesis by
which catalyses the production of the chemical cyclic binding to DNA and acting as a transcription factor
AMP (cAMP) from ATP in the cytoplasm. The cAMP (unit 10). For example the steroid hormone
in turn activates a cascade of enzymes, resulting in oestrogen stimulates the growth of the uterus lining
the breakdown of glycogen to glucose. The cAMP is by being a transcription factor.
called a “second messenger”.
The effect of a hormone is determined not by the hormone itself, but by the receptor in the target cell.
Thus the same hormone can have different effects in different target cells.
Homeostasis
Homeostasis literally means “staying the same” and it refers to the process of keeping the internal body
environment in a steady state. The importance of this cannot be over-stressed, and a great deal of the
hormone system and autonomic nervous system is dedicated to homeostasis.
blood glucose to ensure there is enough glucose available for cellular respiration, but not
concentration enough to lower the blood water potential and dehydrate cells.
blood water potential to prevent loss or gain of water from cells by osmosis.
blood O2 and CO2 to ensure there is sufficient oxygen for respiration and to prevent carbon
concentration dioxide from forming carbonic acid.
blood pressure to maintain blood flow around the body without losing too much fluid in
capillary beds.
blood ion ions, like Na+ , K+ and Ca2+ , are needed for nerve impulses and metabolic
2- 3-
concentration reactions, while others, like CO3 and PO4 are needed as pH buffers.
The problem for the body is that all of these factors will inevitably change due to normal activities like
respiring, eating, exercising and so on. So homeostasis must constantly deal with these changes by making
more changes itself. Homeostasis is therefore described as a dynamic equilibrium because the factors don’t
stay absolutely constant and set in stone, but rather fluctuate slightly over time. In a good homeostatic
system the fluctuations will be very small.
So in a system controlled by negative feedback the level is never maintained perfectly, but constantly
oscillates about the set point. An efficient homeostatic system minimises the size of the oscillations.
We shall look at two examples of homeostasis in detail: temperature and blood glucose.
The thermoregulatory centre is part of the autonomic nervous system, so the various responses are all
involuntary. The exact responses to high and low temperatures are described in the table below:
Erector pili Muscles contract, raising skin hairs and Muscles relax, lowering the skin hairs and
muscles in skin trapping an insulating layer of still, warm allowing air to circulate over the skin,
(attached to skin air next to the skin. Not very effective in encouraging convection and evaporation.
hairs) humans, just causing “goosebumps”.
Brown fat tissue Non-shivering thermogenesis. Increased Decreased respiration in brown fat tissue.
respiration in brown fat tissue –
specialised fat tissue packed with
“uncoupled” mitochondria that respire
triglycerides to produce heat but no ATP.
Especially important in babies and
hibernating mammals.
Adrenal and Glands secrete adrenaline and thyroxine Glands stop releasing adrenaline and
thyroid glands respectively, which increase the thyroxine, so metabolic rate slows.
metabolic rate in different tissues,
especially the liver, so generating heat.
Behaviour Curling up, huddling, finding shelter, Basking, stretching out, finding shade,
putting on more clothes, etc. swimming, removing clothes, etc.
Note that
some of the responses to low temperature actively generate heat (thermogenesis), while others just
conserve heat.
Similarly some of the responses to warm temperatures actively cool the body down, while others just
reduce heat production or transfer heat to the surface.
The body thus has a range of responses available, depending on the internal and external temperatures.
The advantage of being endothermic is that animals can survive in a wide range of environmental
temperatures, and so can colonise almost any habitat, and remain active at night and in cold weather. This
gives endothermic predators an obvious advantage over ectothermic prey. The disadvantage is that it
requires a lot of energy, so endotherms need to eat far more than ectoderms.
The advantage of being ectothermic is that animals use far less energy than endoderms. At rest the
metabolic rate of a reptile is only 10% of that of a mammal of similar size. At night, when the core
temperature of ectotherms drops with the temperature of the surroundings, their metabolic rate drops
still further. This means ectothermic animals need to eat far less than endotherms, and can often survive
for weeks without eating. The disadvantages are that, at certain times of the day, ectotherms can only
move slowly and have slow reactions. This makes them easy prey and poor predators.
The heart rate is controlled by a part of the medulla oblongata called the cardiac (or cardiovascular) centre.
The cardiac centre can send impulses to the SAN of the heart either through sympathetic neurones or
parasympathetic neurones.
Impulses through sympathetic neurones release the Impulses through parasympathetic neurones release
neurotransmitter noradrenaline at the SAN, the neurotransmitter acetylcholine at the SAN,
causing the heart rate to increase. causing the heart rate to decrease.
The cardiac centre can also change the stroke volume by controlling blood pressure. It can increase the
stroke volume by sending nerve impulses to the arterioles to cause vasoconstriction, which increases blood
pressure so more blood fills the heart at diastole. Alternatively it can decrease the stroke volume by
causing vasodilation and reducing the blood pressure.
The overall effects of the cardiac centre in response to exercise are shown in this table:
heart rate stroke volume cardiac output
(beats min-1) (cm3 beat-1) (cm3 min-1)
at rest 75 75 5 600
at exercise 180 120 22 000
Another part of the medulla, called the ventilation centre, controls the breathing rate and causes it to
increase during exercise. Together, these systems provide more oxygen for the muscles to respire.
Response to Stress
The heart rate can also be controlled through the endocrine system, particularly by the hormone
adrenaline. As we’ve already seen, hormonal control is slower but longer-lasting than nervous control, and
hormones can also have more wide-ranging effects. These differences are illustrated by the fight-or-flight
response to stress. In our distant past these stressful situations would be hunting woolly mammoths or
escaping attack by a sabre-tooth tiger, but today the stress is more likely to be running for a bus, playing
sport or even preparing for an exam. Whatever the initial stimulus, the response is coordinated by the
cerebral cortex, which send impulses through the sympathetic system to various organs, especially the
adrenal glands.
The two adrenal glands are located on top of the kidneys. Each gland has an outer layer, called the cortex,
which secretes steroid hormones, and an inner layer, called the medulla (not to be confused with the
medulla oblongata in the brain), which secretes adrenaline. Adrenaline has many target organs:
The SAN responds to adrenaline by increasing the heart rate
Arterioles respond to adrenaline by constricting, which increases the blood pressure and so also
increases the heart stroke volume.
The cardiac centre in the medulla responds by increasing the frequency of sympathetic impulses to the
heart.
Collectively, these responses to adrenaline allow faster respiration and contraction by skeletal muscles. It is
interesting to note that the hormone adrenaline has exactly the same fight-or-flight effects on target organs
as the closely-related neurotransmitter noradrenaline, released from sympathetic neurones.
Excretion
Excretion means the removal of metabolic waste products from cells. There are five important excretory
organs in humans:
• Skin excretes sweat, containing water, ions and urea
• Lungs excrete carbon dioxide and water
• Liver excretes bile, containing bile pigments, cholesterol and mineral ions
• Gut excretes gut mucosa cells, water and bile in faeces. (The bulk of faeces comprises plant fibre
and bacterial cells, which have never been absorbed into the body, so are not excreted but
egested.)
• Kidneys excrete urine, containing urea, mineral ions, water and other toxins from the blood.
This section is mainly concerned with the excretion of nitrogenous waste as urea. The body cannot store
protein in the way it can store carbohydrate and fat, so it cannot keep excess amino acids. The “carbon
skeleton” of the amino acids can be used in respiration, but the nitrogenous amino group must be
excreted.
1. Transamination
In this reaction an amino group is transferred from an amino acid to a keto acid, to form a different amino
acid.
Amino acid 1 + Keto acid 2 → Keto acid 1 + Amino acid 2
In this way scarce amino acids can be made from abundant ones. In adult humans only 11 of the 20 amino
acids can be made by transamination. The others are called essential amino acids, and they must be supplied
in the diet.
2. Deamination
In this reaction an amino group is removed from an amino acid to form ammonia and a keto acid. The most
common example is glutamate deamination:
This reaction is catalysed by the enzyme glutamate dehydrogenase. Most other amino acids are first
transaminated to from glutamate, which is then deaminated. The NADH produced is used in the
respiratory chain; the -ketoglutarate enters the Krebs cycle; and the ammonia is converted to urea (step
3).
3. Urea Synthesis
In this reaction ammonia is converted to urea, ready for excretion by the kidney.
Ammonia is highly toxic, due to the reversal of the glutamate dehydrogenase reaction that would use up all
the -ketoglutarate and so stop the Krebs cycle. Urea is less toxic than ammonia, so it is safer to have in
the bloodstream. The disadvantage is that it “costs” 3 ATP molecules to make one urea molecule. This is
not in fact a single reaction, but is a summary of another cyclic pathway, called the urea (or orthnthine)
cycle.
The Kidney
The kidneys remove urea and other toxic wastes from the blood, forming a dilute solution called urine in
the process. The two kidneys have a very extensive blood supply and the whole blood supply passes
through the kidneys every five minutes, ensuring that waste materials do not build up. The renal artery
carries blood to the kidney, while the renal vein carries blood, now with far lower concentrations of urea
and mineral ions, away from the kidney. The urine formed passes down the ureter to the bladder.
The important part of the kidney is a folded tube called a nephron. There are thousands of nephrons in
each kidney. There are five parts to a nephron, and five steps in producing urine in a nephron:
To understand the process, we start in the ascending limb. The cells of the ascending limb have a special
Na-K-Cl cotransporter protein as well as the usual Na-K-ATPase pump. Together, the proteins act to
pump Na+ , K+ and Cl- ions out of the filtrate into the medulla. This makes the medulla more concentrated
and the filtrate less concentrated. Water would follow by osmosis, but it can’t, because the ascending limb
is impermeable to water. As the filtrate moves up the ascending limb it loses more ions so becomes more
dilute. The ion pumps work at a constant rate, always keeping the medulla a little more concentrated than
the filtrate, so a concentration
gradient is built up in the medulla.
So the countercurrent multiplier ensures there is always a concentration gradient between the filtrate and
the medulla along the whole length of the loop of Henle, enabling the maximum salt gradient to be built up
in the medulla. At the base of the medulla it is three times more concentrated than seawater.
Remember that the collecting duct cells contain aquaporin protein channels that allow water to diffuse very
quickly through the cell membrane. ADH binds to a specific ADH receptor protein in the epithelial cell
membrane, which catalyses the release of the second messenger cAMP (cyclic AMP) inside the endothelial
cells. The cAMP in turn causes vesicles containing aquaporin channels to fuse with the cell membrane thus
making the cell membrane much more permeable to water.
The amount of water in the blood also affects blood pressure. As we saw on p34, blood pressure is
detected by baroreceptors in the aortic and carotid bodies. These baroreceptors also send impulses to the
hypothalamus to control the release of ADH.
This chart shows the volumes of water gained and lost each day by humans and kangaroo rats. Although
the overall daily water turnover per gram of body mass is similar, the desert rat manages to do this without
drinking at all.
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Plant Responses
Like all living organisms, plants can sense and respond to stimuli. For example plants can sense light
direction, gravity and moisture, and can respond by growing in particular direction, flowering or dropping
leaves (abscission). The responses in plants are much slower than those in animals and they usually involve
growth. Plants don’t have a nervous system and their responses are coordinated by plant hormones. Plant
hormones, also known as plant growth substances, are small molecules like animals hormones but with
some significant differences:
Animal hormones are produced in specialised glands, whereas plants don’t have glands and plant
hormones are produced in unspecialised cells in any parts of the plant.
Animal hormones are carried round the whole body in the bloodstream, whereas plant hormones
usually diffuse from cell to cell and have fairly local effects, though they can also be transported through
the xylem and phloem.
Animal hormones affect specific target cells by binding to specific hormone receptor proteins on those
cells, whereas plant hormones generally enter cells and can have many different effects.
Some of the different plant hormones and their effects are summarised in this table:
Plant Hormone Auxin Cytokinin Giberellin Florigen
Chemical structure ?
As this table shows plant hormones generally have multiple effects. They also interact with each other, so
that a particular effect is often the result of contributions from two or more hormones.
When plant hormones promote similar processes, they have a synergistic effect and the overall effect is
enhanced. For example auxin and gibberellin have synergistic actions on shoot growth.
When plant hormones promote opposite processes, they have an antagonistic effect and the overall
effect depends on the balance between the hormones. For example auxin and cytokinin have
antagonistic actions on apical dominance.
Some of the earliest experiments on plant responses investigated this tropism response, and indeed
Charles Darwin and his son Francis studied phototropism in the 1880s. They used coleoptiles – young grass
shoots protected by a smooth cylindrical sheath that grow straight without leaves or buds, so are ideal for
observing growth patterns. The Darwins observed the effects of
light
directional light on these coleoptiles – what we now call phototropism.
Their experiments, and others by later investigators, concluded that a
chemical (a plant hormone named auxin) is synthesized in the coleoptile
tip; asymmetric illumination is detected by
the coleoptile tip and this causes auxin to move into the darker side; auxin
diffuses down the coleoptile; the higher auxin concentration on the darker
side causes more growth on that side; which causes the coleoptile to bend
towards the light source.
In 1934 auxin was identified as a compound called indoleacetic acid, or IAA. IAA is hydrophobic so it can
diffuse through cell membranes and so move around the plant. IAA (auxin) causes growth by stimulating
cell elongation. It activates an enzyme that breaks the bonds between cellulose microfibrils and this loosens
the cell wall and so allows the cell to elongate under the internal turgor pressure.
Cell Division
Tropism work primarily by stimulating cell elongation, but many plant hormones also stimulate cell division.
For example cytokinins are produced by meristem tissue at the tips of roots and shoots where they
stimulate cell division by triggering formation of transcription factors (unit 10). The name cytokinin was
given because the hormone promotes cytokinesis (cytoplasmic division).
Cell Differentiation
As plants grow new cells differentiate into a particular type of plant tissue. Experiments have shown that
differentiation depends on the relative concentrations of two or more plant hormones. For example:
If cytokinin = auxin then cells form an undifferentiated cell mass called a callus.
If cytokinin > auxin then cells differentiate into shoot cells.
If cytokinin < auxin then cells differentiate into root cells.
The two plant hormones are antagonists. This knowledge is very useful in commercial plant tissue culture,
where whole plants are grown from cloned cells.
Apical Dominance
Plants grow from meristem (growing) tissue in buds. Buds are found at the tips, or apex, of shoots and
roots (apical buds) and at branches (lateral buds). Auxin and cytokinin work antagonistically to control bud
growth: auxin inhibits lateral buds while cytokinin stimulates lateral buds. Normally it is important for
plants to grow tall so they can out-compete their neighbours and absorb
more sunlight for photosynthesis, and to help achieve this goal the apical
buds produce auxin, which passes down the phloem and inhibits the
growth of the later buds below. Thus most of the plant’s sugars are used
for growth at the apex. This control is called apical dominance. Further
down the stem the concentration of auxin decreases, so lateral buds can
start to grow, and this explains the “Christmas tree” shape of many
plants, with a wide base and a narrow top.
Seed Germination
A plant seed is in a dormant state to protect the embryo during harsh times and while the seed is
dispersed. When conditions are right the dormancy is broken and the seed germinates, growing a shoot
and root. The plant hormone gibberellin is involved in this germination process.
1. The seed absorbs water. This stimulates the embryo to secrete gibberellin, which diffuses to the outer,
alurone layer.
2. Gibberellin is a transcription factor, and it stimulates the cells of the alurone layer to express the
amylase gene and synthesise the enzyme amylase. The amylase diffuses into the endosperm, where it
digests the starch reserves into the sugars maltose and glucose.
3. The sugars are absorbed by the embryo and used for respiration and as raw materials for biosynthesis
of protein, carbohydrates and lipids to form new cells.
In fact, later experiments showed that the length of darkness is more important than the length of daylight:
So long-day plants are really short-night plants and short-day plants are really long-night plants, but the
original labels are still used. The correct period of darkness is called the photoperiod and the response to
the correct photoperiod is called photoperiodism.
Plants can therefore use the phytochrome system to measure photoperiods. At dawn any Pr is quickly
converted to Pfr, while during the night the Pfr is slowly converted to Pr. The longer the night the more Pfr
will be converted and the lower the Pfr:Pr ratio. Pfr is the physiologically-active form of phytochrome,
though its mode of action is not clear. It appears to be a transcription factor, switching the expression of
various genes.
This signal must therefore be a plant hormone and it has been named florigen. Despite years of research,
florigen has not yet been identified, and it may in fact be a combination of other plant hormones.
Gibberellin appears to act synergistically with florigen to promote flowering.
Other aspects of plant growth and development are also controlled by light, including seed germination, the
size, shape and number of leaves and chlorophyll synthesis. In all cases phytochrome is the light receptor
and it switches on expression on genes. The control of growth and development in plants by specific
wavelengths of light is called photomorphogenesis.