C3.1 Integration of body systems

SL & HL+ AHL sectionsSL and HL, with AHL content · Organisms · Theme C

Big questions

  • How do nerves and hormones work together so the body acts as one coordinated whole?
  • How do feedback loops keep vital variables like heart and breathing rate in check?

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Integration and the hierarchy of subsystems

SL & HLSyllabus understandings: C3.1.1 · C3.1.2

Any system with many parts needs integration: the parts must be coordinated so that together they carry out an overall function. In a car, the engine, gears and brakes must work in step; in a multicellular organism, millions of cells, many tissues and several organ systems must do the same. Integration needs communication between the parts and a way to control what each part does.

A hierarchy of subsystems

Multicellular organisms are organized as a hierarchy of nested subsystems:

  • Cells of the same type, such as muscle fibres, are grouped into tissues.
  • Different tissues are combined into organs, such as a muscle, which contains muscle tissue, connective tissue, blood vessels and nerves.
  • Organs that work together form organ systems, such as the musculoskeletal, nervous, circulatory and respiratory systems.
  • Organ systems together make up the organism.

Each level is a system in its own right, and each is also a component of the level above.

Levels of organization in a cheetahFive boxes linked by arrows: a muscle fibre (cell) is part of skeletal muscle tissue, which forms a leg muscle (organ), which works with bones in the musculoskeletal system (an organ system), which together make up the cheetah (organism). Below, a bar shows that the cheetah's speed is an emergent property that comes from integrating its muscular, skeletal, nervous, respiratory and circulatory systems.Cellmuscle fibreTissueskeletal muscletissueOrgana leg muscleOrgan systemmusculoskeletalsystem(muscles + bones)OrganismcheetahmuscleboneEmergent property: a fast, effective predatorSpeed needs muscles, skeleton, nervous system, lungs, heart and blood working together.No single cell, tissue or organ can sprint: the property appears only from integration.
Levels of organization in a cheetah. Each level is a subsystem of the one above; speed emerges only when the systems are integrated.

Emergent properties

When subsystems are integrated, the whole has emergent properties: abilities that none of the parts has on its own. A cheetah is an effective predator because its body systems work together. Its brain integrates information from its eyes to track prey; its nervous system coordinates powerful contractions of its leg muscles, acting on a long, flexible skeleton; its lungs, heart and blood deliver oxygen and glucose to the muscles and remove CO₂ and heat. No muscle, bone or neuron can sprint at more than 90 km/h, but the integrated animal can.

Nerves, hormones and the blood

SL & HLSyllabus understandings: C3.1.3

Organs in animals are integrated in two ways: by signals, which coordinate what organs do, and by transport of materials and energy between them.

Two signalling systems

Nervous system Endocrine system
Signal Electrical impulses along neurons (chemical at synapses) Hormones (chemical messengers)
Route Along neurons to specific cells Secreted by glands into the blood; carried all round the body
Targets Only the cells the neurons connect to Only cells with receptors for that hormone
Speed Very fast (milliseconds) Slower (seconds to hours)
Duration of effect Usually short-lived Often longer-lasting
Examples Withdrawing the hand from a flame; contracting muscles to run Melatonin and sleep; epinephrine in danger

The two systems work together. The hypothalamus, in the brain, controls much of the endocrine system, and some nerves stimulate glands directly: sympathetic nerves trigger the release of epinephrine from the adrenal glands. Nerves and hormones are covered in more detail in C2.2 Neural signalling and C2.1 Chemical signalling.

Transport by the blood

The blood system links all organs. It carries materials from where they are absorbed or produced to where they are used or removed:

  • oxygen from the lungs to respiring tissues, and CO₂ back to the lungs;
  • glucose, amino acids and other products of digestion from the small intestine to the liver and then to all tissues;
  • urea from the liver, where it is made, to the kidneys, where it is excreted;
  • hormones from endocrine glands to their target cells.

Blood also distributes energy: chemical energy in glucose and fatty acids, and heat from active organs such as the liver and muscles to the skin, where it can be lost. See B3.2 Transport for the structure of blood vessels.

The brain, cerebellum and spinal cord

SL & HLSyllabus understandings: C3.1.4 · C3.1.5 · C3.1.10

The central nervous system (CNS) is the brain and spinal cord. It receives information from receptors, processes it and sends signals to effectors.

The central nervous system: brain and spinal cordA side view of the brain cut down the middle, with the face to the left. The large cerebral hemispheres fill the top. The cerebellum is a smaller, folded structure at the back, below the hemispheres. The brainstem runs down from the centre of the brain; its lowest part, the medulla oblongata, continues into the spinal cord. The hypothalamus is at the base of the brain above the pituitary gland, and the pineal gland is near the centre. Spinal nerves branch from the spinal cord at intervals; each contains sensory and motor fibres.Cerebral hemispheres (cerebrum)integrate information from many inputs;conscious thought, learning, memory;start voluntary movementsPineal glandCerebellumcoordinates skeletal muscle contraction,posture and balanceMedulla oblongata (brainstem)heart rate, ventilation rate, swallowingSpinal nervesbundles of sensory and motor fibresSpinal cordintegrates unconscious processes (reflexes);carries signals to and from the brainHypothalamuslinks nervous andendocrine systemsPituitary glandfront of head ←
The brain cut down the middle (face to the left), the spinal cord and spinal nerves.

The brain as an integrating organ

The cerebral hemispheres (cerebrum) are the main centre for integrating information. They receive impulses from many sensory inputs at once and combine them. When you catch a ball, for example, information from your eyes about the ball’s movement is combined with information about the position of your arm and hand and with memories of earlier catches, and the brain then sends a coordinated set of signals to your muscles. The brain also stores information as memories, and learning happens when experience changes the connections between neurons, so a later response can be different.

Conscious and unconscious processes

  • Conscious processes are ones we are aware of and can control, such as deciding to pick up a cup. They are coordinated in the cerebral hemispheres.
  • Unconscious processes happen without our awareness or decision, such as a withdrawal reflex, the knee-jerk reflex, or the control of heart rate.

The spinal cord acts as an integrating centre for unconscious processes. Reflexes are processed in its grey matter (cell bodies and synapses of relay and motor neurons), while its outer white matter consists of myelinated axons carrying impulses up to and down from the brain.

The cerebellum

The cerebellum is at the back of the brain, below the cerebral hemispheres. It coordinates skeletal muscle contraction and balance. It receives information about the movement the brain intends to make, together with feedback from stretch receptors in muscles and joints, from the balance organs of the inner ear and from the eyes. It compares what is happening with what was intended and fine-tunes the timing and force of muscle contractions, so movements are smooth and accurate and posture is maintained. Skills such as riding a bicycle become automatic as the cerebellum learns them. Damage to the cerebellum causes clumsy, jerky movements and an unsteady walk, although the muscles themselves are not weakened.

Sensory neurons, motor neurons and nerves

SL & HLSyllabus understandings: C3.1.6 · C3.1.7 · C3.1.8

Input: sensory neurons

Receptors detect stimuli. Some are specialized receptor cells, such as the rods and cones of the eye; others are the endings of the sensory neurons themselves, such as the free nerve endings that detect pain. Sensory neurons carry nerve impulses from receptors into the CNS: through the dorsal roots of spinal nerves into the spinal cord, or through cranial nerves directly into the brain. From the spinal cord the information is relayed up to the cerebral hemispheres, where sensations are perceived. A sensory neuron’s cell body lies in a ganglion just outside the CNS.

Output: motor neurons

To make a voluntary movement, the cerebral hemispheres send impulses down the spinal cord, where they are passed to motor neurons. A motor neuron’s cell body is in the grey matter of the spinal cord (or brainstem); its long axon runs out through a ventral root and a nerve to a skeletal muscle. At the motor end plate the motor neuron releases a neurotransmitter that stimulates the muscle fibres to contract.

Nerves

A nerve is a bundle of many nerve fibres. Most nerves are mixed: they contain the fibres of both sensory neurons (carrying impulses towards the CNS) and motor neurons (carrying impulses away from it). In a transverse section you can see:

  • an outer protective sheath of connective tissue (the epineurium);
  • several bundles (fascicles) of fibres, each wrapped in its own sheath (the perineurium);
  • myelinated fibres: axons surrounded by a myelin sheath, which looks like a ring with a dark rim and a pale centre;
  • unmyelinated fibres: smaller axons, often grouped within a Schwann cell, with no myelin ring;
  • small blood vessels in the connective tissue.
Transverse section of a nerveA round nerve cut across. An outer protective sheath of connective tissue (the epineurium) surrounds four bundles of nerve fibres, each wrapped in its own sheath (the perineurium). Small blood vessels run in the connective tissue between the bundles. Inside each bundle are many nerve fibres cut across: myelinated fibres look like rings, with a dark myelin sheath around a pale axon, and unmyelinated fibres are smaller axons grouped together inside Schwann cells without a myelin sheath. A nerve contains fibres of both sensory and motor neurons.Epineuriumprotective sheath of connective tissuearound the whole nervePerineuriumsheath around each bundle (fascicle)Blood vesselMyelinated fibreaxon (pale) in a myelin sheath (dark ring)Unmyelinated fibressmall axons in a Schwann cell, no myelinSensory and motor fibres runtogether in the same nerve.
Transverse section of a nerve: a protective sheath around bundles of myelinated and unmyelinated fibres.

Myelinated fibres conduct impulses much faster than unmyelinated ones (see C2.2 Neural signalling).

The pain reflex arc

SL & HLSyllabus understandings: C3.1.9

A reflex is a rapid, automatic, involuntary response to a stimulus. The pathway the impulses follow is a reflex arc. The pain withdrawal reflex protects the body from damage, and the IB example has skeletal muscle as the effector.

The pain reflex arc
brainfree nerve endings(pain receptors)stimulus: heatspinal nerve: sensory + motor fibresdorsal root ganglion(sensory cell body)dorsal rootrelayneurongrey matterwhite mattermotor neuroncell bodyventral rootbicepsmuscle(effector)to the brainin white mattersensory neuronrelay neuronmotor neuronneuron to brain
Text description of this animation

On the left an arm, with the upper arm vertical and the forearm held forwards so that the fingertip is just above a candle flame. On the right the spinal cord is shown in transverse section, dorsal side at the top: a butterfly-shaped area of grey matter surrounded by white matter, with a central canal. A spinal nerve from the arm divides into a dorsal root, which has a swelling (the dorsal root ganglion) containing the cell body of the sensory neuron, and a ventral root. Inside the grey matter a single relay neuron connects the sensory neuron to a motor neuron whose cell body is in the ventral horn. The motor neuron's axon runs out through the ventral root and the spinal nerve to the biceps. A separate neuron carries a signal up the white matter to the brain. The six steps follow a nerve impulse around the arc and show the arm flexing to pull the hand away.

  1. Stimulus at the receptor. The fingertip touches a flame. Free nerve endings of a sensory neuron in the skin act as pain receptors — heat that could damage the tissue stimulates them, and nerve impulses start. No separate receptor cell is needed.
  2. Sensory neuron to the spinal cord. Impulses travel along the long axon of the sensory neuron, through a spinal nerve and the dorsal root, into the grey matter of the spinal cord. The sensory neuron's cell body lies in the dorsal root ganglion, a swelling on the dorsal root.
  3. Relay neuron in the grey matter. In the grey matter the sensory neuron synapses with a single relay neuron (interneuron), which passes the signal on to a motor neuron. This is where the reflex is integrated, without any input from the brain.
  4. Motor neuron to the muscle. The relay neuron synapses with a motor neuron whose cell body is in the ventral horn of the grey matter. Impulses travel out along its axon, through the ventral root and the same spinal nerve, to the arm.
  5. The effector responds. At motor end plates the motor neuron stimulates the biceps, a skeletal muscle (the effector), to contract. The arm flexes at the elbow and the hand is pulled away from the flame. The response is involuntary and fast because the brain is not involved.
  6. The brain becomes aware. The sensory neuron also synapses with a neuron that carries impulses up the spinal cord, in the white matter, to the brain. The pain is felt consciously only when these signals reach the cerebral hemispheres — after the hand has already started to move.

The five parts

  1. Receptor: free nerve endings of a sensory neuron in the skin of the hand act as pain receptors. Heat or pressure strong enough to damage tissue stimulates them, and nerve impulses begin.
  2. Sensory neuron: carries the impulses along its axon, in a spinal nerve, through the dorsal root into the spinal cord. Its cell body is in the dorsal root ganglion.
  3. Relay neuron (interneuron): lies entirely within the grey matter of the spinal cord. The sensory neuron synapses with it, and it synapses with the motor neuron. In this example there is a single relay neuron.
  4. Motor neuron: its cell body is in the ventral part of the grey matter; its axon leaves the cord through the ventral root and runs in the same spinal nerve back to the arm.
  5. Effector: the motor neuron stimulates a skeletal muscle, here the biceps, to contract. The arm bends at the elbow and the hand is pulled away.

Why it is fast and involuntary

Only three neurons and two synapses in the spinal cord are involved, and the response does not wait for the brain. The sensory neuron also passes impulses to neurons that carry them up the white matter to the brain, so a moment later the pain is felt. The brain can then decide what to do next, and in some cases (for example holding on to a hot plate to avoid dropping it) can override the reflex.

Hormones: the hypothalamus, melatonin and epinephrine

SL & HLSyllabus understandings: C3.1.11 · C3.1.12 · C3.1.13

Hormonal control by the hypothalamus, pineal and adrenal glands
pineal glandhypothalamuspituitary glandthyroid glandheartadrenal glandskidneysHypothalamus and pituitary glandhypothalamusneurosecretory cellsanteriorpituitaryposteriorpituitaryportalvesselsTSH, FSH, LH, GH…ADH, oxytocinblood carries hormones to target organsthyroxine: negative feedbackMelatonin from the pineal glanddarklightlight12:0018:0000:0006:0012:00melatonin in bloodtime of daylight → retina → SCN in the hypothalamus → pineal glandlight suppresses secretion; darkness allows itEpinephrine (adrenaline) from the adrenal glandsHeart rate and stroke volume increaseBronchioles widen; ventilation rate risesLiver breaks down glycogen to glucoseArterioles to skeletal muscle widenArterioles to gut and skin narrowFatty acids released from adipose tissuePupils dilate; alertness increasesMore O₂ and glucose reach muscles,ready for intense contraction
Text description of this animation

On the left, an outline of the human body showing the pineal gland, hypothalamus and pituitary gland in the head, the thyroid gland in the neck, the heart, and the adrenal glands on top of the kidneys. On the right, a panel changes with each step. The first four steps show the hypothalamus above the pituitary gland: neurosecretory cells send hormones down their axons to the posterior pituitary, or release releasing hormones into portal blood vessels that run to the anterior pituitary, which then secretes its own hormones into the blood. A negative feedback arrow shows thyroxine inhibiting the hypothalamus and pituitary. The fifth step shows a 24-hour graph of melatonin in the blood: low during the day, rising after dark, peaking in the middle of the night and falling before morning. The last step lists the effects of epinephrine from the adrenal glands.

  1. The hypothalamus controls the pituitary. The hypothalamus, at the base of the brain, receives nerve impulses from many brain regions and monitors the blood. It links the nervous and endocrine systems by controlling the pituitary gland just below it, whose hormones regulate many other glands.
  2. Hormones made in the hypothalamus. Neurosecretory cells in the hypothalamus make ADH and oxytocin. The hormones pass down the cells' axons to the posterior pituitary, which stores them and releases them into the blood when impulses arrive.
  3. Releasing hormones and the anterior pituitary. Other neurosecretory cells secrete releasing hormones into portal blood vessels that lead to the anterior pituitary. In response, its cells secrete their own hormones, such as TSH, FSH, LH and growth hormone, into the blood.
  4. Target glands and negative feedback. TSH stimulates the thyroid gland to secrete thyroxine. Rising thyroxine inhibits the release of TRH by the hypothalamus and of TSH by the anterior pituitary, keeping thyroxine within a narrow range (negative feedback).
  5. Melatonin and the sleep–wake cycle. Light detected by the retina inhibits melatonin secretion by the pineal gland, via the suprachiasmatic nucleus (SCN) of the hypothalamus. After dark secretion rises, peaking in the middle of the night; melatonin promotes sleep, and levels fall again before waking.
  6. Epinephrine prepares the body for action. In danger, stress or vigorous exercise, sympathetic nerves stimulate the adrenal glands to secrete epinephrine (adrenaline) into the blood. It acts on many tissues at once, so skeletal muscles receive more oxygen and glucose for intense contraction.

The hypothalamus and pituitary gland

The hypothalamus, at the base of the brain, links the nervous and endocrine systems. It receives impulses from other parts of the brain and monitors the blood, and it controls the pituitary gland, which hangs just below it. Some hormones are made by neurosecretory cells in the hypothalamus and released from the pituitary; others are made by the pituitary itself in response to releasing hormones from the hypothalamus. Many pituitary hormones control other endocrine glands: for example TSH stimulates the thyroid gland, and FSH and LH act on the ovaries and testes. When the hormone from the target gland rises, it inhibits the hypothalamus and pituitary: negative feedback keeps hormone levels stable. (You need only this general account, not the differences between the anterior and posterior pituitary.)

Melatonin and the sleep–wake cycle

The pineal gland secretes melatonin in a diurnal (daily) pattern. Light detected by the retina inhibits secretion, through the suprachiasmatic nucleus (SCN) in the hypothalamus, which acts as the body’s master clock. So melatonin concentration is low during the day, rises in the evening as it gets dark, peaks in the middle of the night and falls towards morning. High melatonin promotes sleepiness and lowers body temperature, helping to establish a regular circadian rhythm of sleeping and waking of about 24 hours. Jet lag happens because the melatonin rhythm takes several days to shift to a new time zone, and bright screens late in the evening can delay melatonin secretion.

Epinephrine and vigorous activity

In danger, stress or intense exercise, sympathetic nerves stimulate the adrenal glands, on top of the kidneys, to secrete epinephrine (adrenaline) into the blood. It has widespread effects, which together prepare the body for vigorous activity (“fight or flight”):

  • heart rate and stroke volume increase, so more blood reaches the muscles;
  • bronchioles widen and ventilation rate increases, so more oxygen is absorbed;
  • the liver breaks down glycogen and releases glucose into the blood;
  • arterioles supplying skeletal muscles widen, while those supplying the gut and skin narrow;
  • pupils dilate and alertness increases.

The result is a greater supply of oxygen and glucose to skeletal muscle, so it can respire faster and contract more intensely.

Feedback control of heart rate

SL & HLSyllabus understandings: C3.1.14

Heart rate is controlled by negative feedback: a change is detected and a response is made that reverses it. The sinoatrial (SA) node sets the heartbeat (see B3.2 Transport), but the cardiovascular centre in the medulla oblongata changes its rate.

The receptors and where they are

  • Baroreceptors detect blood pressure by the stretch of the artery wall. They are in the walls of the aortic arch and the carotid sinuses (swellings of the carotid arteries in the neck).
  • Chemoreceptors detect blood pH and the concentrations of CO₂ and O₂. They are in the carotid bodies and aortic bodies, small structures close to the baroreceptors; there are also chemoreceptors in the medulla.

The response

The medulla sends impulses along two kinds of nerve to the SA node:

  • sympathetic (cardiac accelerator) nerves increase heart rate. They also act on the ventricular muscle, increasing the force of contraction, so stroke volume rises;
  • the vagus nerve (parasympathetic) decreases heart rate.
Change detected Signals from the medulla Effect
Blood pressure rises More vagus, less sympathetic Heart rate and stroke volume fall; pressure falls back
Blood pressure falls (e.g. standing up quickly) More sympathetic, less vagus Heart rate and stroke volume rise; pressure is restored
CO₂ rises, pH falls (exercise) More sympathetic, less vagus Heart rate and stroke volume rise; blood flow to muscles increases
Simulation Feedback control of heart rate and ventilation rate

Heart rate

Heart rate
70 beats min⁻¹
Stroke volume
70 cm³
Cardiac output
4.9 dm³ min⁻¹

Ventilation rate

Blood CO₂ (pCO₂)
5.3 kPa
Blood pH
7.39
Ventilation rate
12 breaths min⁻¹
Tidal volume
0.5 dm³
Heart rate and ventilation rate against exercise intensity02040608010004080120160200Exercise intensity (% of maximum) →Rate per minute →heart rate (beats)ventilation rate (breaths)

Heart rate Ventilation rate

At rest: blood pH 7.39, heart rate 70 beats per minute and ventilation rate 12 breaths per minute. The vagus nerve dominates, keeping the heart slower than the SA node's own rate.

Show data table
Exercise (%)Blood pHHeart rate (min⁻¹)Ventilation rate (min⁻¹)
07.397012
207.399318
407.3811623
607.3713929
807.3616234
1007.3418540

Qualitative model: values are realistic in size but simplified. In real exercise, signals from the motor cortex and from receptors in muscles and joints also raise heart and ventilation rates, and adrenaline adds to the sympathetic effect.

Text description of this simulation

Two signal pathways and a graph. In the heart-rate panel, chemoreceptors in the carotid and aortic bodies (sensing blood pH, CO₂ and O₂) and baroreceptors in the carotid sinus and aortic arch (sensing blood pressure) send impulses to the cardiovascular centre in the medulla. The medulla sends impulses along a sympathetic nerve, which speeds the heart and strengthens its contractions, and along the vagus nerve (parasympathetic), which slows it; both act on the sinoatrial node, and the sympathetic nerve also acts on the ventricular muscle, increasing the force of contraction (stroke volume). In the ventilation panel, chemoreceptors in the brainstem and in the carotid and aortic bodies signal to the respiratory centre in the medulla, which controls the diaphragm and intercostal muscles. Raising exercise intensity increases CO₂ production and lowers blood pH, so heart rate, stroke volume, ventilation rate and tidal volume all rise, which keeps pH close to normal. Raising blood pressure increases baroreceptor firing and vagus activity, so heart rate falls. A thought-experiment switch blocks the chemoreceptor signals to the respiratory centre: ventilation then cannot rise, CO₂ accumulates and pH falls far below normal. The model is qualitative.

Feedback control of ventilation rate

SL & HLSyllabus understandings: C3.1.15

Why blood pH changes

Blood pH is normally kept between about 7.35 and 7.45. The main cause of change is carbon dioxide. CO₂ from cell respiration dissolves in the plasma and, mostly inside red blood cells (where carbonic anhydrase speeds up the reaction), reacts with water to form carbonic acid, which dissociates:

CO₂ + H₂O ⇌ H₂CO₃ ⇌ H⁺ + HCO₃⁻

So when respiration speeds up, for example during exercise, more CO₂ enters the blood, more H⁺ is released and pH falls. In very intense exercise, lactic acid from anaerobic respiration lowers pH further. When CO₂ is removed faster than it is produced, pH rises.

The control loop

  1. Chemoreceptors in the brainstem (medulla) detect the fall in pH; chemoreceptors in the carotid and aortic bodies detect pH, CO₂ and O₂ in arterial blood.
  2. They send impulses to the respiratory centre in the medulla.
  3. The respiratory centre sends impulses more often along nerves to the diaphragm (the phrenic nerves) and the external intercostal muscles, which contract more often and more strongly.
  4. Ventilation rate and tidal volume (depth) increase, so CO₂ is removed from the blood faster.
  5. Blood CO₂ falls and pH rises back towards normal. When pH returns to normal, impulses from the chemoreceptors decrease and ventilation slows: negative feedback.

In the simulation above, tick the thought-experiment box to block the chemoreceptor signals. Ventilation can no longer rise, and in hard exercise CO₂ builds up and pH falls far outside the normal range: a dangerous acidosis.

Nervous control of peristalsis

SL & HLSyllabus understandings: C3.1.16

Peristalsis is the wave of muscle contraction that moves food along the gut. The gut wall has two layers of smooth muscle: an inner circular layer and an outer longitudinal layer. Behind the bolus of food the circular muscle contracts, narrowing the gut and squeezing the food forwards; in front of it the circular muscle relaxes, so the gut is wider there.

Nervous control of swallowing, peristalsis and egestionTop: the gut from mouth to anus in three zones. Swallowing is started voluntarily by the central nervous system; peristalsis along the oesophagus, stomach and intestines is involuntary and coordinated by the enteric nervous system; egestion of faeces through the anus is under voluntary control by the central nervous system. Bottom: a section of gut wall with a bolus of food. The circular muscle is contracted behind the bolus, narrowing the gut and pushing the bolus forwards, and relaxed in front of it. The longitudinal muscle lies outside the circular muscle, and a network of enteric neurons lies between the two layers.Swallowingstarted voluntarily:brain (CNS)Peristalsisoesophagus → stomach → intestines:involuntary, coordinated by the entericnervous system (ENS) in the gut wallEgestionvoluntary control ofthe anus: CNSmouth → anusPeristalsis in a section of gut wallbolus movescircular muscle contractedbehind the boluscircular muscle relaxedin front of the boluslongitudinal muscleenteric neurons
Swallowing and egestion are under voluntary control by the CNS; peristalsis in between is involuntary, coordinated by the enteric nervous system.

Two levels of control

  • Swallowing is started voluntarily. The tongue (skeletal muscle under the control of the brain) pushes food to the back of the mouth. From there, swallowing continues automatically and the food enters the oesophagus.
  • Peristalsis is involuntary. From the oesophagus, through the stomach and the small and large intestines, waves of contraction are coordinated by the enteric nervous system (ENS): networks of neurons within the gut wall, between and beneath the muscle layers. The ENS detects stretching of the gut wall by food and coordinates contraction behind the bolus and relaxation in front of it, so material passes along in the right direction and at a suitable rate. A section of gut removed from the body can still carry out peristalsis, showing the ENS can work without the brain (though the CNS can speed it up or slow it down).
  • Egestion is under voluntary control. Faeces collect in the rectum. The anus is closed by sphincter muscles, one of which is skeletal muscle that the CNS can keep contracted until it is convenient to egest. Babies do not yet have this control; it is learned in early childhood.

Phototropism and observing tropic responses

AHLSyllabus understandings: C3.1.17 · C3.1.18

Plants cannot move from place to place, but they can change their direction of growth. A tropism is a directional growth response to a stimulus that comes from one direction. In positive phototropism, shoots grow towards light coming from one side (lateral, or unilateral, light). This brings the leaves into brighter light for photosynthesis. Only phototropism of shoots is required.

Observing tropisms in seedlings

Young seedlings, such as cress, radish or oat seedlings, respond within hours, so they are ideal for experiments. They can be grown in a light-proof box with a slit on one side, with a control group given light evenly from above.

Observing and measuring phototropism in seedlingsThree panels. First, seedlings grow in a light-proof box with a slit on one side, so light reaches them from one direction only; a control group receives light evenly from above. Second, the seedlings are recorded qualitatively with simple drawings at the start, when they are straight, and after 24 hours, when their shoots have bent towards the light. Third, the response is measured quantitatively: a vertical line is drawn up from where the shoot starts to bend, a second line along the tip, and the angle between them, the angle of curvature, is measured with a protractor, here 45 degrees.1 Set-up2 Record (qualitative)3 Measure (quantitative)light-proof box with a slit:light from one side only(control: light from above)0 h24 hlightsimple line drawings, to scale,with time and light directionθ = 45°angle between the vertical andthe tip = angle of curvature
Seedlings in unilateral light: setting up, recording qualitatively with drawings, and measuring the angle of curvature.
  • Qualitative data are descriptions: careful, simple line drawings of the seedlings at the start and after a set time, with the light direction marked, and notes such as “all shoots bent towards the slit”.
  • Quantitative data are measurements: the angle of curvature, between the vertical and the direction of the shoot tip, measured with a protractor, often on a photograph.

Precision, accuracy and reliability

  • Precision is limited by the measuring instrument and method: a protractor marked in 1° divisions can only be read to about ±1°, and it is hard to judge exactly where a curved shoot’s tip points. Photographing seedlings against a grid and measuring with image software can improve precision.
  • Accuracy (closeness to the true value) can be improved by photographing from directly side-on, defining the measurement lines in the same way every time and calibrating against a known angle.
  • Reliability comes from repeats: measuring many seedlings, calculating a mean and a measure of spread, and keeping other variables (light intensity, distance from the lamp, temperature, seedling age) the same.

Phytohormones and auxin efflux carriers

AHLSyllabus understandings: C3.1.19 · C3.1.20

Phytohormones

Plants use a variety of chemicals as phytohormones (plant hormones). They are signalling molecules produced in one part of the plant that control growth, development and responses to stimuli, often in other parts of the plant, and they are effective at very low concentrations. Examples:

Phytohormone Some effects
Auxin Cell elongation in shoots; phototropism; apical dominance; lateral root formation
Cytokinins Cell division; growth of side buds
Gibberellins Stem elongation; seed germination
Abscisic acid Closing stomata in drought; seed dormancy
Ethylene (ethene) Fruit ripening; leaf fall

The effect of a phytohormone often depends on its concentration, on the tissue and on the other phytohormones present.

How auxin gradients are made

The main natural auxin is indole-3-acetic acid (IAA). Its distribution in the plant is controlled by auxin efflux carriers (PIN proteins):

  1. The cell wall is acidic, so some of the auxin there is uncharged (IAAH). Uncharged auxin can diffuse into a cell across the plasma membrane.
  2. The cytoplasm is less acidic (about pH 7), so auxin loses H⁺ and becomes a negative ion (IAA⁻). Charged auxin cannot diffuse through the membrane, so it is trapped.
  3. IAA⁻ can leave the cell only through auxin efflux carriers, and a cell can place these on just one side of its plasma membrane.
  4. If every cell in a tissue places its carriers on the same side, auxin is passed from cell to cell in one direction and becomes concentrated in one part of the plant.
Auxin efflux carriers move auxin from cell to cellThree plant cells in a row, separated by cell walls. Auxin enters each cell by diffusing across the plasma membrane in its uncharged form. The cytoplasm is less acidic than the cell wall, so inside the cell auxin loses a hydrogen ion and becomes negatively charged, which traps it. Auxin efflux carriers are located only in the right-hand plasma membrane of every cell, so auxin leaves each cell on the right, gains a hydrogen ion in the acidic cell wall and diffuses into the next cell. Because every cell places its carriers on the same side, auxin is moved in one direction through the tissue and builds up at one end. Proton pumps in the membrane keep the cell wall acidic.IAA⁻ trapped(cytoplasm pH ≈ 7)IAA⁻ trapped(cytoplasm pH ≈ 7)IAA⁻ trapped(cytoplasm pH ≈ 7)cell wallpH ≈ 5.5IAA⁻ + H⁺ → IAAH,which diffuses into the next cellproton pump keepsthe wall acidicauxin efflux carrier (PIN protein), on one side of each cell onlyIAA⁻ carried out by efflux carriersuncharged IAAH diffuses innet (polar) movement of auxin
Polar transport of auxin: each cell has efflux carriers on the same side, so auxin moves from cell to cell in one direction.

The process depends on proton pumps, which use ATP to keep the cell wall acidic, so the whole transport system is active. Cells can move their efflux carriers to a different side of the cell, which redirects the flow of auxin — this is what happens when a shoot is lit from one side.

How auxin makes cells elongate

AHLSyllabus understandings: C3.1.21

Auxin promotes the elongation of shoot cells. A plant cell is surrounded by a cell wall of cellulose microfibrils, held together by cross-links with other wall polysaccharides. The cell can only grow larger if the wall is loosened.

Acid growth

  1. Auxin binds to receptors and stimulates proton pumps in the plasma membrane, which use ATP to pump H⁺ ions out of the cytoplasm into the cell wall (the apoplast).
  2. The cell wall becomes acidic.
  3. The low pH activates proteins called expansins, which loosen the cross-links between cellulose molecules.
  4. The cell’s turgor pressure stretches the loosened wall, water enters by osmosis and the cell elongates.

From auxin gradients to phototropism

When a shoot is lit from one side:

  1. Phototropins (photoreceptors) in the shoot tip detect the direction of the light.
  2. Auxin efflux carriers are moved to the sides of the cells facing away from the light, so auxin is transported to the shaded side.
  3. The shaded side has the higher auxin concentration, so its cells elongate faster than those on the lit side.
  4. Unequal elongation makes the shoot bend towards the light.
How auxin causes phototropismAHL
light from abovelight from one sideshoot tipmakes auxinblue lightdetected byphototropinsangle ofcurvature≈ 45°auxinCells below the shoot tipauxin pumped towards the shaded sidemore auxin → more elongationshaded sidelit sideauxin efflux carrier (PIN protein)Acid growth in a shaded-side cellcell wallcytoplasmH⁺H⁺H⁺H⁺H⁺H⁺H⁺ATP1 Auxin activatesproton pumps2 H⁺ pumped into thewall, which becomes acidic3 Low pH activatesexpansins: cross-linksbetween cellulose loosen4 Turgor stretches thewall: the cell elongatescellulosecross-linkexpansin
Text description of this animation

On the left, a seedling growing in a pot. On the right, a close-up of a block of cells just below the shoot tip, in four columns: the two left columns are on the side away from the light (the shaded side) and the two right columns are on the lit side. Auxin is shown as orange dots. With light from above, auxin is spread evenly and the shoot grows straight. With light from the right, auxin efflux carriers (PIN proteins) move to the membranes facing the shaded side, and auxin is pumped across to the shaded side. A close-up of a cell wall shows acid growth: proton pumps secrete hydrogen ions into the wall, expansins loosen the cross-links between cellulose microfibrils and the cell elongates. Finally the shaded-side cells are longer than the lit-side cells and the shoot bends towards the light; the angle of curvature, about 45 degrees, is marked.

  1. Light from above. With light from above, the shoot tip makes auxin, which is transported down both sides of the shoot equally. Cells on both sides elongate at the same rate, so the shoot grows straight up.
  2. Light from one side. Light now comes from the right only. Photoreceptor proteins (phototropins) in the shoot tip absorb blue light and detect the direction it comes from.
  3. Auxin moves to the shaded side. Auxin efflux carriers (PIN proteins) are moved to the membranes facing the shaded side of the cells, so auxin is pumped sideways from cell to cell. Auxin becomes more concentrated on the shaded side.
  4. Acid growth in shaded-side cells. Auxin activates proton pumps that secrete H⁺ into the cell wall (apoplast). The acidic wall activates expansins, which loosen the cross-links between cellulose microfibrils, so turgor pressure stretches the wall and the cell elongates.
  5. The shoot bends towards the light. Shaded-side cells elongate more than lit-side cells, so the shoot curves towards the light (positive phototropism). The angle of curvature, between the vertical and the tip, can be measured to compare responses.

Auxin and cytokinin coordinate roots and shoots

AHLSyllabus understandings: C3.1.22

A plant needs its shoot system (for photosynthesis) and its root system (for water and mineral ions) to stay in proportion. Two phytohormones, travelling in opposite directions, help to integrate their growth:

  • Shoot tips (and young leaves) produce auxin, which is transported down the plant to the roots. There it promotes the formation of lateral roots, extending the root system.
  • Root tips produce cytokinin, which is carried up to the shoots in the xylem. There it promotes cell division and the growth of side buds, extending the shoot system.
Auxin and cytokinin coordinate shoot and root growthA young plant with its shoot above the soil and roots below. Auxin is made in the shoot tip and young leaves and is transported down the plant to the roots, where it promotes the formation of lateral roots; in the shoot it inhibits the growth of side buds. Cytokinin is made in the root tips and is carried up to the shoot in the xylem, where it promotes cell division and the growth of side buds. Because each organ system sends a signal that promotes growth of the other, the sizes of the root system and the shoot system stay matched.soilauxincytokininshoot tiproot tipAuxinmade in shoot tips and young leavestransported down to the rootsroots: promotes lateral root formationshoot: inhibits side buds (apical dominance)Cytokininmade in root tipscarried up to the shoots in the xylemshoot: promotes cell division and side budsIntegrationA large root system sends more cytokininup, so the shoot grows more; a large shootsends more auxin down, so more roots form.The balance (ratio) of the two hormoneskeeps root and shoot growth matched.
Shoot tips send auxin to the roots and root tips send cytokinin to the shoots, so each promotes the growth of the other.

How they interact

  • If the root system grows, more cytokinin reaches the shoot and the shoot grows more; a larger shoot sends more auxin to the roots, so more roots form. Growth of each part is therefore linked to the size of the other.
  • In the shoot the two hormones have opposite effects on side buds: auxin from the apical bud inhibits their growth (apical dominance), while cytokinin promotes it. The balance between them decides how bushy the shoot becomes.
  • In plant tissue culture, a high ratio of auxin to cytokinin makes a mass of cells form roots, and a high ratio of cytokinin to auxin makes it form shoots, which shows that the ratio of the hormones matters, not just their amounts.

Ethylene and positive feedback in fruit ripening

AHLSyllabus understandings: C3.1.23

Ethylene (IUPAC name ethene, C₂H₄) is a gaseous phytohormone. It triggers the changes of ripening in fruits such as tomatoes, bananas and apples:

  • starch is converted to sugars, so the fruit becomes sweeter;
  • enzymes break down cell walls, so the fruit softens;
  • chlorophyll is broken down and other pigments are made, so the colour changes;
  • scent compounds are released.
Positive feedback in fruit ripeningA loop of two boxes. Ethylene (ethene) stimulates the changes of ripening, and ripening tissue makes more ethylene, so each increases the other: positive feedback. The changes listed are starch converted to sugars, cell walls softened, chlorophyll broken down, new pigments made and scents released. On the right, a ripening fruit gives off ethylene gas, which diffuses to neighbouring fruits and triggers their ripening too, so a whole group of fruits ripens rapidly and at the same time.Ethylene (ethene)a gaseous phytohormoneRipeningchanges in the fruit++positivefeedbackethylene → ripening → more ethylene → …Ripening changesstarch → sugars (sweeter)cell walls broken down (softer)chlorophyll broken downnew pigments (colour)scent compounds releasedSynchronized ripeningethylene gas diffusesto nearby fruitsfruits ripen quickly, together
Ethylene stimulates ripening and ripening stimulates ethylene production: positive feedback. Ethylene gas spreads to neighbouring fruits.

Positive feedback

Ripening tissue produces more ethylene, and more ethylene speeds up ripening. Because the response amplifies the original stimulus, this is positive feedback (compare the negative feedback that keeps heart rate and blood pH stable). The result:

  • Rapid ripening: once ripening starts, it accelerates and is completed quickly.
  • Synchronized ripening: ethylene is a gas, so it diffuses from a ripening fruit to the fruits around it and triggers their ripening too. All the fruits on a plant ripen at about the same time.

This benefits the plant: a large crop of ripe, attractive fruit at once draws animals that eat the fruit and disperse the seeds. People use the same mechanism: bananas are picked unripe, transported, then ripened in rooms containing ethylene, and “one ripe apple in a bag” speeds the ripening of the others. Positive feedback is useful here because it drives a process to completion; for comparison with homeostasis see D3.3 Homeostasis.

Must-know definitions

System integration and emergent properties
Integration is the coordination of the parts of a system so that together they perform an overall function. Cells, tissues, organs and organ systems are nested subsystems, and their integration gives the organism emergent properties that no part has alone, such as a cheetah's ability to hunt at high speed.
Nervous vs endocrine signalling
The nervous system sends electrical impulses along neurons to specific target cells: fast and short-lived. The endocrine system secretes hormones into the blood, which reaches every tissue, but only cells with the right receptors respond: slower and often longer-lasting.
Sensory and motor neurons
A sensory neuron carries impulses from receptors to the central nervous system (spinal cord and brain). A motor neuron carries impulses from the central nervous system to an effector, such as a skeletal muscle, which it stimulates to contract.
Nerve
A bundle of nerve fibres (the axons of many neurons), usually of both sensory and motor neurons, surrounded by a protective sheath of connective tissue. It contains both myelinated and unmyelinated fibres.
Pain reflex arc
The pathway of a rapid, involuntary response: free nerve endings (pain receptors) → sensory neuron → relay neuron in the grey matter of the spinal cord → motor neuron → skeletal muscle (effector). The brain is informed but does not take part.
Baroreceptors and chemoreceptors
Baroreceptors in the walls of the aortic arch and carotid sinuses detect blood pressure. Chemoreceptors in the carotid and aortic bodies detect blood pH and concentrations of CO2 and O2; chemoreceptors in the medulla (brainstem) detect pH changes caused by CO2.
Medulla oblongata
The lowest part of the brainstem. Its cardiovascular centre adjusts heart rate and stroke volume through sympathetic and vagus nerves, and its respiratory centre adjusts ventilation rate through nerves to the diaphragm and intercostal muscles.
Enteric nervous system (ENS)
A network of neurons in the wall of the gut that coordinates peristalsis involuntarily, without needing input from the brain. Only swallowing (at the start) and egestion (at the end) are under voluntary control by the CNS.
Auxin efflux carrierAHL
A membrane protein (a PIN protein) that moves the charged form of auxin out of a cell. Auxin can diffuse into cells but cannot leave without these carriers, so placing them on the same side of every cell moves auxin in one direction through a tissue.
Acid growthAHL
Auxin stimulates proton pumps that secrete H+ into the cell wall (apoplast). The low pH activates expansins, which loosen the cross-links between cellulose molecules, so turgor pressure can stretch the wall and the cell elongates.
Positive feedback in fruit ripeningAHL
Ethylene (ethene) stimulates ripening, and ripening tissue produces more ethylene. Because the response amplifies the stimulus, ripening speeds up, and ethylene gas spreading between fruits makes them ripen together.

Common exam mistakes

  • Loses marks Hormones travel along nerves to their target organs.

    Write instead Hormones are secreted by endocrine glands into the blood and carried all round the body. Nerves carry electrical impulses along neurons; the two systems use different routes.

  • Loses marks In a pain reflex the brain decides to move the hand away.

    Write instead The reflex is integrated in the grey matter of the spinal cord by a relay neuron. Signals reach the brain too, but pain is felt only after the muscle has already started to contract.

  • Loses marks The cell body of the sensory neuron is in the grey matter of the spinal cord.

    Write instead The sensory neuron's cell body is in the dorsal root ganglion, outside the grey matter. The cell bodies of relay neurons and motor neurons are in the grey matter, and the motor axon leaves through the ventral root.

  • Loses marks Baroreceptors detect carbon dioxide and chemoreceptors detect blood pressure.

    Write instead Baroreceptors detect pressure (stretch of the artery wall); chemoreceptors detect chemical changes: pH, CO2 and O2.

  • Loses marks The vagus nerve speeds up the heart and the sympathetic nerve slows it down.

    Write instead It is the other way round: sympathetic (accelerator) nerves increase heart rate and the force of contraction; the vagus nerve (parasympathetic) decreases heart rate. Both act on the SA node, and sympathetic nerves also act on the ventricular muscle, increasing the force of contraction (stroke volume).

  • Loses marks During exercise breathing speeds up because chemoreceptors detect that blood oxygen has fallen.

    Write instead The main stimulus is CO2: more CO2 from respiration forms carbonic acid, which releases H+ and lowers blood pH. Chemoreceptors detect the fall in pH. Arterial O2 changes little in normal exercise.

  • Loses marks Peristalsis is a voluntary action controlled by the brain.

    Write instead Peristalsis is involuntary and coordinated by the enteric nervous system in the gut wall. Only the start of swallowing and egestion are voluntary (CNS).

  • Loses marks Light destroys auxin on the lit side of a shoot, or makes the lit side produce more auxin.

    Write instead Unilateral light causes auxin to be redistributed to the shaded side by auxin efflux carriers, so the shaded side has the higher concentration.

    AHL
  • Loses marks Auxin makes the cells on the shaded side divide faster.

    Write instead In phototropism auxin causes cell elongation, by acidifying the cell wall (acid growth), not faster cell division.

    AHL
  • Loses marks Ethylene production in ripening fruit is controlled by negative feedback.

    Write instead It is positive feedback: ripening increases ethylene production, which speeds ripening further, so ripening is rapid and synchronized.

    AHL

Self-test

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Multiple-choice questions

  1. Question 1Which statement correctly distinguishes hormonal signalling from nervous signalling in animals?
  2. Question 2In a pain withdrawal reflex arc involving the spinal cord, where is the cell body of the sensory neuron?
  3. Question 3A person's blood pressure suddenly rises. Which sequence describes the response that restores normal blood pressure?
  4. Question 4During a sprint, an athlete's ventilation rate rises. Which change in the blood is mainly responsible for stimulating the chemoreceptors?
  5. Question 5In a light micrograph of a transverse section of a nerve, many fibres appear as rings with a dark outer rim and a pale centre, while others appear as small dots with no rim. What do the rings and the dots show?
  6. Question 6Nurses working night shifts often sleep poorly during the day. Blood samples show that their melatonin concentration stays low while they are working under bright lights at night. Which explanation is best supported?
  7. Question 7AHLA student measures the angle of curvature of 20 seedlings after 24 hours in light from one side, using a protractor marked in 1° divisions. Which change would most improve the reliability of the mean angle?
  8. Question 8AHLWhich statement explains how auxin can be moved in one direction through a plant tissue?
  9. Question 9AHLShoot segments were treated with auxin and with an inhibitor of the plasma-membrane proton pumps. Compared with segments treated with auxin alone, the segments with the inhibitor elongated very little. What is the most likely reason?
  10. Question 10AHLUnripe bananas stored in a sealed room ripen much faster if a few ripe bananas are added. What is the explanation?

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Prompt

What is an emergent property? Give the C3.1 example.

Last reviewed 19 October 2026. Spotted a mistake? Report it.

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