B3.2 Transport
SL & HL+ AHL sectionsSL and HL, with AHL content · Organisms · Theme B
Big questions
- How do animals and plants move fluids around large bodies using pressure?
- In what ways are blood vessels and xylem alike, and how are they different?
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Capillaries, arteries and veins
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Text description of this visual
The first model shows three blood vessels side by side, each cut across so the front face shows the wall in transverse section. The artery has a thick wall compared with its narrow lumen: a thin tunica intima whose lining looks folded, a thick tunica media of circular smooth muscle with wavy layers of elastic fibres, and an outer tunica externa of collagen-rich connective tissue. The vein has a thin wall and a wide, slightly flattened lumen; its far end is cut open lengthways to show a pocket valve whose cusps point towards the heart. The capillary, drawn far larger than to scale, has a wall one endothelial cell thick on a thin basement membrane, a narrow pore between two endothelial cells, fenestrations through the cells and red blood cells in single file. The second model (higher level) is a mammalian heart cut open from top to bottom and seen from the front, so the heart's right side is on the viewer's left. It shows the four chambers, the thick-walled left ventricle, the septum, the tricuspid and bicuspid valves with their tendons, the semilunar valves, the venae cavae, pulmonary artery, pulmonary veins and aorta, coronary arteries on the surface and the sinoatrial node, and it steps through the route blood takes through both sides of the heart.
Artery, vein and capillary
- Tunica externa
- The outer coat of tough connective tissue rich in collagen fibres. Collagen has high tensile strength, so it stops the artery bulging outwards (an aneurysm) or bursting when blood pressure is high.
- Tunica media (thick)
- The thickest layer of an artery wall, made of circular smooth muscle and elastic fibres. Its thickness lets the artery withstand the high pressure of blood pumped from the heart.
- Elastic fibres
- Layers of elastin in the tunica media. They are stretched by each pulse of blood (storing energy) and recoil between beats, pushing blood onwards, which keeps the pressure up and evens out the flow.
- Smooth muscle cells
- Smooth muscle cells (nuclei shown dark) run in circles around the artery. Contraction narrows the lumen (vasoconstriction) and relaxation widens it (vasodilation), adjusting blood flow.
- Tunica intima (endothelium)
- A single layer of smooth, flat endothelial cells that lines the lumen and reduces friction. In micrographs of arteries it often looks folded (corrugated), because the elastic layer beneath it recoils.
- Lumen of artery (narrow)
- The lumen is narrow compared with the thickness of the wall, which helps to maintain high blood pressure. In sections it is usually round.
- Thin wall of vein
- The tunica media is thin, with few muscle and elastic fibres, because blood pressure in veins is low and there is no pulse. The thin, flexible wall can be squeezed flat by surrounding skeletal muscles.
- Lumen of vein (wide)
- Wide compared with the thickness of the wall, which gives little resistance to the slow, low-pressure flow of blood back to the heart. Veins often look flattened or irregular in sections.
- Pocket valve
- Pocket-shaped flaps (cusps) attached to the vein wall. Blood flowing towards the heart pushes them back against the wall; blood starting to flow backwards fills the pockets, which meet and close the vein.
- Capillary wall one cell thick
- A single layer of very thin endothelial cells, so substances diffuse only a very short distance between the blood and the tissues. (This capillary is drawn far larger than to scale.)
- Pore between endothelial cells
- Narrow gaps between neighbouring endothelial cells let water and small solutes leak out of the blood to form tissue fluid, while blood cells and most plasma proteins stay inside.
- Basement membrane
- A thin layer of extracellular fibrous proteins around the endothelium. It supports the capillary and acts as a filter, letting small and medium-sized particles through but not large proteins.
- Fenestrations
- Larger pores through the endothelial cells themselves, found in fenestrated capillaries where exchange must be especially rapid, such as the glomerulus of the kidney, the villi of the small intestine and endocrine glands.
- Red blood cells in single file
- The lumen of a capillary (about 5–10 µm) is only just wide enough for red blood cells to pass in single file. Capillaries branch so much that their total surface area for exchange is huge.
Heart, frontal section (AHL)
- Right atrium
- A thin-walled collecting chamber that receives deoxygenated blood from the body through the venae cavae. Its contraction tops up the right ventricle through the tricuspid valve.
- Left atrium
- Receives oxygenated blood from the lungs through the pulmonary veins and pumps it into the left ventricle. Atria need only thin walls because they pump blood a short distance at low pressure.
- Right ventricle
- Pumps deoxygenated blood to the lungs through the pulmonary artery. Its wall is thinner than the left ventricle's because the lungs are close and their capillaries would be damaged by high pressure.
- Left ventricle (thickest wall)
- Its thick wall of cardiac muscle contracts strongly enough to produce the high pressure (about 120 mmHg) needed to pump oxygenated blood through the aorta to all the organs of the body.
- Septum
- A thick wall between the left and right sides of the heart. It keeps oxygenated and deoxygenated blood apart, so the heart works as two pumps side by side (a double pump).
- Tricuspid valve (right AV valve)
- The atrioventricular valve between the right atrium and right ventricle, with three flaps. It closes when the ventricle contracts, preventing backflow; tendons (chordae tendineae) stop it turning inside out.
- Bicuspid (mitral) valve (left AV valve)
- The atrioventricular valve between the left atrium and left ventricle, with two flaps. It closes as soon as ventricular pressure rises above atrial pressure, so blood cannot flow back into the atrium.
- Semilunar valves
- The aortic and pulmonary valves, each with three pocket-shaped cusps (two show in this cut). They open when ventricular pressure exceeds the pressure in the artery and close to stop blood flowing back.
- Venae cavae
- The superior and inferior venae cavae return deoxygenated blood at low pressure from the upper and lower parts of the body to the right atrium.
- Pulmonary artery
- Carries deoxygenated blood from the right ventricle to the lungs. Its trunk lies in front of the aorta, so it is drawn crossing over it before dividing into the left and right pulmonary arteries.
- Pulmonary veins
- Four pulmonary veins (two from each lung) bring oxygenated blood to the left atrium. They open into the back wall of the atrium, so in this model they come from behind the heart.
- Aorta
- The largest artery. It receives oxygenated blood at high pressure from the left ventricle; its first branches are the coronary arteries, and branches from its arch supply the head and arms.
- Coronary arteries
- Branch from the aorta just above the aortic valve and run over the surface of the heart, supplying its cardiac muscle with oxygen and glucose. A blockage starves the muscle beyond it of oxygen.
- Sinoatrial node (pacemaker)
- A small group of specialized cardiac muscle cells in the wall of the right atrium, near where the superior vena cava enters. It initiates each heartbeat with an electrical impulse that spreads through the atria.
- Venae cavae to right atrium. Deoxygenated blood from the body returns at low pressure through the superior and inferior venae cavae and fills the right atrium. Meanwhile the ventricles are contracting, so the tricuspid valve is closed.
- Right atrium to right ventricle. As the ventricles relax, pressure in the right ventricle falls below that in the right atrium. The tricuspid valve opens and blood flows into the ventricle, and atrial contraction tops it up. The pulmonary valve is closed.
- Right ventricle to pulmonary artery. The right ventricle contracts. Rising pressure closes the tricuspid valve and opens the pulmonary semilunar valve, and deoxygenated blood is pumped into the pulmonary artery, which divides to carry it to both lungs.
- Pulmonary veins to left atrium. Oxygenated blood returns from the lungs through the four pulmonary veins into the left atrium. The bicuspid valve is closed because the ventricles are contracting.
- Left atrium to left ventricle. As the left ventricle relaxes, the bicuspid (mitral) valve opens and blood flows from the left atrium into the left ventricle; atrial contraction completes the filling. The aortic valve is closed.
- Left ventricle to aorta. The thick-walled left ventricle contracts strongly. The bicuspid valve closes, the aortic semilunar valve opens and oxygenated blood is pumped at high pressure into the aorta. Both sides of the heart contract at the same time.
Blood flows from the heart through arteries, which branch into narrow arterioles, then through capillaries, and back to the heart through venules and veins. Each type of vessel has a wall suited to its job.
Capillaries: adapted for exchange
Capillaries are where materials are exchanged between the blood and the tissues (or, in the lungs and gut, with the outside world). They have three key adaptations:
- Large surface area. Capillaries branch and rejoin repeatedly to form dense capillary networks, so their total surface area is enormous and every active cell is close to one. Their narrow lumen (about 5–10 µm) means red blood cells pass in single file, close to the wall.
- Thin, permeable wall. The wall is a single layer of very thin endothelial cells on a basement membrane, so the diffusion distance is tiny. Small gaps (pores) between the cells let water and small solutes leak out to form tissue fluid, while blood cells and most plasma proteins stay in.
- Fenestrations. Fenestrated capillaries also have larger pores through the endothelial cells themselves. They are found where exchange must be especially rapid, such as the glomerulus of the kidney (filtration of blood), the villi of the small intestine (absorption) and endocrine glands (release of hormones).
Telling arteries from veins in micrographs
| Feature | Artery | Vein |
|---|---|---|
| Wall thickness relative to lumen | thick wall, narrow lumen | thin wall, wide lumen |
| Tunica media | thick: much smooth muscle and elastic tissue | thin: little muscle or elastic tissue |
| Shape in section | usually round, holds its shape | often flattened or irregular |
| Lining | often looks folded (corrugated) | smooth |
| Valves | none (except at the exit from the heart) | pocket valves in many veins |
Arteries withstand and maintain high pressure
Each time the ventricles contract (systole), a surge of blood at high pressure enters the arteries; between beats (diastole) the heart pumps nothing into them. Artery walls are built to cope with this.
- Collagen fibres in the tunica externa (and media) are very strong in tension, so the artery does not bulge outwards (an aneurysm) or burst under the highest pressures.
- Elastic fibres (elastin) in the tunica media are stretched by each surge, storing potential energy. When the ventricles relax, the fibres recoil, squeezing the blood onwards. This keeps arterial pressure from falling to zero between beats (it stays around 80 mmHg in large arteries) and makes the flow more continuous.
- Smooth muscle in the tunica media is arranged in circles. Its contraction narrows the lumen (vasoconstriction); relaxation widens it (vasodilation). Arterioles have the most muscle relative to their size, so they control how much blood each tissue receives.
- A thick wall and a relatively narrow lumen help to maintain high pressure; a smooth endothelium in the tunica intima reduces friction.
The stretch and recoil of the artery wall with every heartbeat is what you feel as the pulse. By the time blood reaches the capillaries, its pressure has fallen and the pulse has smoothed out.
Measuring pulse rate
There is one pulse for every beat of the heart, so the pulse rate equals the heart rate. You can feel it wherever an artery runs close to the skin over something firm:
- the radial pulse, on the palm side of the wrist below the thumb;
- the carotid pulse, in the neck in the soft groove beside the windpipe (press gently and on one side only).
Method
- Rest quietly for a few minutes first if you want a resting rate.
- Place the tips of your index and middle fingers on the artery. Do not use your thumb, which has a pulse of its own.
- Count the beats for a fixed time and convert to beats per minute (bpm): ×4 for 15 s, ×2 for 30 s, ×1 for 60 s.
- Repeat and calculate a mean.
A resting adult heart rate is usually about 60–100 bpm.
Time left: — · Beats counted: 0
Pulse oximeter: — bpm
Choose a counting time, press Start, then tap once for every pulse you see. Your count × 4 gives beats per minute for a 15 s count.
Show results table
| Count time | Beats | × | Heart rate (bpm) | Digital (bpm) |
|---|---|---|---|---|
| No counts yet. | ||||
Simulated pulse at a steady rate. Real heart rate varies slightly from beat to beat, and a pulse oximeter averages over several beats.
Text description of this simulation
A drawing of a wrist with two fingertips on the radial pulse. After you press Start, the artery glows each time a pulse of blood passes and you tap a button once for every beat. When the time is up, the count is multiplied up to one minute: by 4 for a 15 second count, by 2 for 30 seconds. The result is compared with a pulse oximeter reading. A short count is quicker but less precise, because each beat missed or added changes a 15 second result by 4 beats per minute.
Traditional versus digital methods
Digital methods include pulse oximeters, which clip on a fingertip, shine light through it and detect the change in light absorption as each pulse of blood arrives; fitness watches, which use a similar optical sensor; and chest-strap monitors, which detect the heart’s electrical activity. They are quick, give continuous readings and avoid counting errors, but they can be thrown off by movement, cold fingers or a loose fit. Counting by hand needs no equipment but depends on the counter. A short count is quicker, but each missed or extra beat changes a 15 s result by 4 bpm.
Veins return blood to the heart
By the time blood has passed through capillaries, its pressure is low and steady: there is no pulse in veins. Veins are adapted to get this slow-moving blood back to the heart.
- Valves. Many veins, especially in the limbs, contain pocket valves: flaps of tissue attached to the wall like pockets. Blood flowing towards the heart pushes the flaps against the wall, so the valve is open. If blood starts to flow backwards, it fills the pockets, which meet in the middle and close the vein. Blood can therefore only move one way, towards the heart.
- Thin, flexible walls. Veins have a thin tunica media with little muscle or elastic tissue. When nearby skeletal muscles contract, they bulge and press on the veins, squeezing them flatter and pushing blood along; the valves make sure it goes towards the heart. This is sometimes called the skeletal muscle pump, and it is one reason why walking helps prevent blood pooling in the legs.
- Wide lumen. A large lumen gives little resistance to flow, so the low pressure is enough to move a large volume of blood. At any one time most of the blood in the body is in the veins.
Coronary artery occlusion and heart disease
Cardiac muscle works without stopping, so it needs a constant supply of oxygen and glucose for aerobic respiration. This comes not from the blood inside the heart’s chambers but from the coronary arteries, the first branches of the aorta, which run over the surface of the heart.
Causes
In atherosclerosis, fatty deposits called plaques (atheroma), made of lipids including cholesterol, fibrous tissue and white blood cells, build up in the walls of arteries just beneath the endothelium.
- The plaque narrows the lumen and makes the artery wall less elastic, so blood flow is reduced.
- If the surface of a plaque ruptures, a blood clot (thrombus) forms there. The clot can block the artery completely: occlusion.
Consequences
- A narrowed coronary artery may supply enough blood at rest but not during exercise, when cardiac muscle needs more oxygen. The result is chest pain called angina and shortness of breath on exertion.
- If a coronary artery is occluded, the cardiac muscle it supplies receives no oxygen. It cannot respire aerobically, stops contracting and begins to die: a myocardial infarction (heart attack). The heart may beat irregularly or stop.
Together these conditions are called coronary heart disease (CHD).
Lumen area left (assuming even narrowing): 49%. At rest the artery supplies 1.0 of a demand of 1: the heart muscle gets enough oxygen.
Qualitative model in arbitrary units (resting demand = 1). The plaque is drawn on one side, as it often is, but the area left is calculated as if the lumen narrowed evenly all round. Real narrowings vary, and symptoms depend on which artery is affected and on other vessels that may supply the same muscle.
Text description of this simulation
A coronary artery in cross-section beside bars showing the oxygen supplied to the heart muscle it serves and the muscle's demand. As the plaque (atheroma) grows, the lumen narrows. At rest the supply is maintained until the narrowing is severe, but during exercise the narrowed artery cannot deliver enough, so the muscle is short of oxygen (angina). If the plaque ruptures and a blood clot forms, the artery is blocked completely (occlusion): the muscle it supplies receives no oxygen and starts to die, a myocardial infarction. The model is qualitative.
Evaluating epidemiological data
Epidemiology studies the patterns and causes of disease in populations. It has identified risk factors for CHD, including high blood pressure, smoking, high blood cholesterol (especially LDL), obesity, lack of exercise, diabetes, older age, being male and a family history of CHD.
A correlation coefficient (r) quantifies how strongly two variables are linearly related. It runs from −1 (perfect negative correlation) through 0 (no linear correlation) to +1 (perfect positive correlation).
- A low or zero correlation coefficient is evidence against a hypothesis that one variable affects the other.
- A high correlation coefficient supports a hypothesis but does not prove causation. For example, countries where people eat more saturated fat tend to have higher CHD rates, but those countries may also differ in smoking, exercise, wealth or other foods. These confounding variables could explain the pattern.
Water transport from roots to leaves
Plants move large volumes of water from the roots to the leaves in xylem vessels, with dissolved mineral ions. Most of this water is lost from the leaves by transpiration. Remarkably, the movement is entirely passive.
- Evaporation. In the leaf, water evaporates from the moist cell walls of the spongy mesophyll cells into the air spaces. The water vapour then diffuses out through the stomata into the drier air outside (see B3.1 Gas exchange).
- Capillary action in cell walls. Cellulose in cell walls is hydrophilic. The water lost is replaced by water drawn through the narrow spaces between the cellulose fibres, by adhesion of water to cellulose and cohesion between water molecules. This water is pulled from the nearest xylem vessels in the leaf veins.
- Tension in the xylem. Pulling water out of the xylem puts the water inside under tension (a negative pressure). Because of cohesion (hydrogen bonding between water molecules, see A1.1 Water), the whole column of water in the xylem is pulled upwards as a continuous thread, from the roots to the leaves. Adhesion to the vessel walls also helps.
- Water enters the roots. As water is pulled up, it is replaced by water moving across the root from the soil.
This is the transpiration pull, or cohesion–tension mechanism. The energy comes from the heat that causes evaporation, ultimately from the Sun; no ATP is used to move water, and the vessels themselves are dead.
Reduced motion is on: use the arrows to step through. Change this in Display settings.
Text description of this animation
A single xylem vessel runs from a root (bottom) through the stem to a vein in a leaf (top). The leaf is shown in section with palisade and spongy mesophyll, air spaces and a stoma between two guard cells. Water evaporates from moist spongy mesophyll cell walls and the vapour diffuses out through the stoma. Water drawn through the cell walls to replace it is pulled out of the xylem, putting the water column under tension; cohesion between water molecules transmits the pull down to the roots, where water enters from the soil. A final higher-level step shows root pressure, when root cells pump mineral ions into the xylem and water follows by osmosis.
- Evaporation in the leaf. Water evaporates from the moist cell walls of spongy mesophyll cells into the air spaces inside the leaf, which become saturated with water vapour. Heat is the energy source that drives evaporation.
- Vapour diffuses out. Water vapour diffuses out of the leaf through open stomata, from the humid air spaces into the drier air outside. This loss of water vapour from the leaf is transpiration.
- Water drawn through cell walls. Evaporated water is replaced by water drawn through the narrow spaces between cellulose fibres in the cell walls, by capillary action (adhesion to hydrophilic cellulose and cohesion between water molecules). This water is pulled out of the xylem in the leaf vein.
- Tension in the xylem. Drawing water out of the xylem leaves the water inside under tension (negative pressure). Cohesion — hydrogen bonds between water molecules — transmits the pull down the continuous column to the roots. Lignified walls stop the vessel collapsing.
- Water enters the roots. As water is pulled up the xylem, water moves into the root xylem from the surrounding cells, and soil water enters root hair cells by osmosis. All of this is passive; the xylem vessels are dead and the energy comes from evaporation in the leaves.
- Root pressure (AHL). When transpiration is too slow, as in very humid air or in spring before deciduous trees open their leaves, root cells actively pump mineral ions into the xylem. Water follows by osmosis, creating a positive pressure (root pressure) that pushes the sap upwards.
Xylem vessels are adapted for water transport
Xylem vessels develop from columns of cells arranged end to end. As they mature, the cells die and their structure is modified:
- No cell contents. The plasma membranes, cytoplasm and nuclei break down, leaving an empty tube. Nothing obstructs the flow, and the movement of water must be passive.
- No end walls (or incomplete ones). The end walls between the cells are removed, so each vessel is a long continuous tube in which flow of xylem sap is unimpeded.
- Lignified walls. The side walls are thickened and impregnated with lignin, a strong, rigid polymer. Under tension, water pulls inwards on the walls; lignin stops the vessels collapsing. In young xylem the thickening is laid down as rings or a spiral; in older xylem it covers most of the wall.
- Pits. Gaps in the lignified thickening, where only a thin layer of cellulose wall remains, let water move sideways into and out of the vessel, for example into neighbouring vessels or out to the cells of a leaf.
Lignified xylem also gives the plant mechanical support: wood is mostly xylem.
Tissues in dicot stems and roots
A plan diagram shows the outlines of tissues in an organ, without drawing individual cells. You need to draw plan diagrams of transverse sections (TS) of the stem and root of a dicotyledonous plant from micrographs, and annotate them with the functions of the tissues.
Stem
In a young dicot stem, the vascular bundles are arranged in a ring near the outside. In each bundle, phloem is on the outer side and xylem on the inner side, separated by a thin layer of cambium. Outside the ring is the cortex, covered by the epidermis; inside the ring is the pith.
Root
In a dicot root the vascular tissue is in the centre. The xylem forms a star or cross shape, with groups of phloem between its arms. The vascular tissue is surrounded by a ring of endodermis, then a wide cortex and an outer epidermis, many of whose cells grow root hairs.
Tissue fluid: formation, exchange and reuptake
Body cells are not bathed by blood. They are surrounded by tissue fluid, which forms from blood plasma in the capillaries.
Release and reuptake
- Arteriole end. Blood arrives from the arterioles at relatively high pressure (roughly 35 mmHg). This forces plasma out through the pores of the capillary wall: pressure filtration. Water and small solutes leave; blood cells and most plasma proteins are too large and stay in the blood.
- Venule end. Resistance in the capillary reduces the blood pressure (to roughly 15 mmHg). The plasma proteins that stayed in the blood give it a higher solute concentration than the tissue fluid, which tends to draw water back in by osmosis. Once the blood pressure has fallen this low, tissue fluid can drain back into the capillary.
Exchange with cells
As tissue fluid flows between cells, substances are exchanged across the plasma membranes by diffusion, facilitated diffusion and active transport. Cells take up oxygen, glucose, amino acids, fatty acids and ions, and release carbon dioxide, urea and other waste products. The fluid that drains back into the capillaries is therefore lower in oxygen and nutrients and higher in wastes, which the blood carries to the lungs, liver and kidneys.
| Blood plasma | Tissue fluid | |
|---|---|---|
| Water, ions, glucose, amino acids | present | present at similar concentrations |
| Plasma proteins (e.g. albumin, fibrinogen) | high concentration | very little |
| Red blood cells and platelets | carried in blood | absent |
| Oxygen and nutrients | higher (arriving) | lower (being used by cells) |
| Carbon dioxide and urea | lower | higher (released by cells) |
Excess tissue fluid drains into lymph ducts
More fluid leaves the capillaries than drains back into them, and if the excess stayed in the tissues they would swell (oedema). It is removed by the lymphatic system.
- Tissues contain blind-ended lymph capillaries with thin walls that have gaps between their cells, so tissue fluid (and any protein that has leaked out) can enter easily. Once inside, the fluid is called lymph.
- Lymph capillaries join into larger lymph ducts (lymph vessels), which contain valves that prevent backflow. Like veins, they rely on surrounding muscles squeezing them, and the valves make sure the lymph moves in one direction.
- The largest lymph ducts empty the lymph into veins near the heart (the subclavian veins), so the fluid is returned to the blood circulation.
In this way a few litres of fluid a day return to the blood that would otherwise accumulate in the tissues.
Single and double circulation
In a single circulation blood passes through the heart once for each complete circuit of the body; in a double circulation it passes through twice.
Bony fish: single circulation
The fish heart has one atrium and one ventricle. The ventricle pumps deoxygenated blood to the gills, where it is oxygenated, and the blood then flows straight on to the other body organs before returning to the heart. Passing through the gill capillaries lowers the pressure a lot, so blood flows on to the body organs at low pressure and more slowly.
Mammals: double circulation
The mammalian heart is a double pump with two separate sides:
- Pulmonary circulation: the right ventricle pumps deoxygenated blood through the pulmonary artery to the lungs, and oxygenated blood returns through the pulmonary veins to the left atrium. The pressure is relatively low, which protects the delicate capillaries of the alveoli.
- Systemic circulation: the left ventricle pumps oxygenated blood through the aorta at high pressure to all the other organs, and deoxygenated blood returns through the venae cavae to the right atrium.
Returning blood to the heart after the lungs lets it be pumped again at high pressure, so oxygen reaches the tissues quickly. This supports the high metabolic rate of mammals, and high pressure is needed for processes such as filtration in the kidneys (see D3.3 Homeostasis).
The mammalian heart
The heart is a muscular pump that delivers blood under pressure to the arteries. Switch the 3D model at the top of the page to Heart, frontal section to explore it and to step through the route blood takes.
| Structure | Adaptation for pumping |
|---|---|
| Cardiac muscle | contracts and relaxes rhythmically throughout life without fatigue; contraction is myogenic (it starts within the muscle, not from a nerve); branched cells are linked so impulses spread quickly (see B2.3 Cell specialization) |
| Pacemaker (sinoatrial node) | a group of specialized muscle cells in the wall of the right atrium that initiates each heartbeat |
| Atria | thin-walled collecting chambers that fill with blood from the veins and pump it a short distance into the ventricles |
| Ventricles | thick muscular walls generate high pressure to pump blood into the arteries; the left ventricle wall is thicker than the right, because it pumps to the whole body rather than the nearby lungs |
| Atrioventricular valves | the tricuspid (right) and bicuspid or mitral (left) valves prevent backflow into the atria when the ventricles contract; tendons stop them inverting |
| Semilunar valves | the pulmonary and aortic valves prevent backflow from the arteries into the ventricles when the ventricles relax |
| Septum | separates the left and right sides, so oxygenated and deoxygenated blood do not mix and each side pumps at its own pressure |
| Coronary vessels | coronary arteries carry oxygenated blood from the aorta to the cardiac muscle; coronary veins drain it back to the right atrium |
Tracing the flow of blood
Venae cavae → right atrium → tricuspid valve → right ventricle → pulmonary semilunar valve → pulmonary artery → lungs → pulmonary veins → left atrium → bicuspid valve → left ventricle → aortic semilunar valve → aorta.
The cardiac cycle
The cardiac cycle is the sequence of events in one heartbeat. At 75 beats per minute it lasts 60 ÷ 75 = 0.8 s. Contraction is called systole and relaxation diastole. The two sides of the heart work at the same time; the description below follows the left side.
- Initiation. The sinoatrial node (SAN, the pacemaker) sends out an electrical impulse that spreads through the walls of both atria.
- Atrial systole. The atria contract, pushing blood through the open atrioventricular valves to finish filling the ventricles. The semilunar valves are closed.
- Ventricular systole. After a short delay (the impulse passes to the ventricles through the atrioventricular node), the ventricles contract, starting from the apex (bottom) and spreading upwards, which pushes blood towards the arteries. Ventricular pressure rises above atrial pressure, so the AV valves close (the first heart sound). When ventricular pressure exceeds aortic pressure, the semilunar valves open and blood is ejected.
- Diastole. The ventricles relax. When their pressure falls below aortic pressure the semilunar valves close (the second heart sound). When it falls below atrial pressure the AV valves open, and blood that has collected in the atria flows into the ventricles. The aorta’s elastic recoil keeps blood flowing.
Reduced motion is on: use the arrows to step through. Change this in Display settings.
Text description of this animation
The left atrium, left ventricle and aorta (with the right side of the heart faded behind) next to a graph of pressure against time for one heartbeat at 75 beats per minute (0.8 seconds). The graph shows aortic pressure between about 80 and 120 mmHg, left ventricular pressure rising from near zero to about 120 mmHg and falling again, and a low atrial pressure. A cursor moves along the graph while panels show whether the bicuspid (atrioventricular) and aortic (semilunar) valves are open or closed. Each valve opens or closes exactly where two pressure curves cross. The animation is slowed down.
- Atrial systole. The sinoatrial node fires and the impulse spreads through the atrial walls, so the atria contract. Atrial pressure is slightly higher than ventricular pressure, so blood flows through the open bicuspid valve and the ventricle finishes filling. The aortic valve is closed.
- Ventricular systole begins. After a short delay the ventricle contracts. As soon as its pressure rises above atrial pressure the bicuspid valve closes (the first heart sound). With both valves closed, pressure rises steeply but no blood leaves yet.
- Blood is ejected. When ventricular pressure exceeds aortic pressure (about 80 mmHg) the aortic semilunar valve opens and blood is ejected into the aorta. Both pressures rise to a peak of about 120 mmHg, the systolic pressure. The atrium relaxes and starts to refill.
- Diastole begins. The ventricle relaxes. When its pressure falls below aortic pressure the semilunar valve closes (the second heart sound), making a small notch in the aortic curve. Both valves are closed while ventricular pressure falls steeply.
- Ventricle fills. When ventricular pressure falls below atrial pressure the bicuspid valve opens and blood that collected in the atrium flows into the ventricle. Elastic recoil of the aorta keeps blood moving as aortic pressure falls slowly to about 80 mmHg, the diastolic pressure.
Interpreting blood pressure
Blood pressure is usually measured in an artery of the upper arm and written as systolic/diastolic, for example 120/80 mmHg.
- Systolic pressure is the maximum arterial pressure, reached during ventricular systole while the semilunar valves are open.
- Diastolic pressure is the minimum arterial pressure, reached at the end of diastole, just before the semilunar valves open again.
A resting reading persistently at or above about 140/90 mmHg is usually classed as high blood pressure (hypertension), a risk factor for coronary heart disease.
Root pressure
Transpiration pull only works when water is evaporating from the leaves. Sometimes a plant needs to move water and minerals up the xylem when there is little or no transpiration:
- when the air is very humid, so little water evaporates;
- in spring, before the leaves of deciduous plants have opened, when xylem vessels must be refilled with sap and minerals delivered to the growing buds.
In these conditions plants generate root pressure:
- Cells in the root, next to the xylem, actively transport mineral ions (such as K⁺ and nitrate) into the xylem vessels, using ATP from respiration.
- The solute concentration of the xylem sap rises (its water potential falls; see D2.3 Water potential).
- Water follows by osmosis from the surrounding root cells into the xylem.
- With water entering a confined space, a positive pressure builds up that pushes the sap upwards.
Root pressure is positive, whereas transpiration produces tension (negative pressure). It is usually much smaller than the tensions generated by transpiration. Its effects can be seen when sap oozes from the cut stump of a plant, or when droplets of water appear at leaf edges on a humid morning (guttation).
Step 6 of the transpiration animation above shows root pressure.
Phloem: sieve tubes and companion cells
Translocation is the movement of carbon compounds, mainly sucrose, in phloem sap from sources to sinks.
- Sources release carbon compounds: photosynthesising leaves and green stems, and storage organs when their stores are being mobilised.
- Sinks use or store them: roots, growing shoots, developing fruits and seeds, and storage organs that are filling up.
Sieve tube elements
Phloem sieve tubes are columns of living cells, sieve tube elements, joined end to end.
- Sieve plates: the end walls between elements are perforated by large pores, so sap flows easily from one element to the next.
- Reduced cytoplasm and organelles, and no nucleus: almost the whole cell is open to the flow of sap.
- A plasma membrane is kept, so the cell stays alive and can control what enters and leaves.
Companion cells
Each sieve tube element has one or more companion cells beside it.
- They have a nucleus, which supports the sieve tube element, and many mitochondria, which make the ATP for active transport of sucrose.
- Many plasmodesmata (narrow cytoplasmic channels through the shared wall) connect each companion cell to its sieve tube element, so sucrose can pass between them.
How the adaptations work together
- Loading at a source. Companion cells use ATP to pump sucrose in by active transport; it passes through plasmodesmata into the sieve tube.
- The high solute concentration draws water in from nearby xylem by osmosis, raising the hydrostatic pressure in the sieve tube.
- Unloading at a sink. Sucrose is removed from the sieve tube for use or storage, water follows it out by osmosis and the pressure falls.
- Sap flows along the pressure gradient from source to sink. Different sieve tubes can carry sap in different directions, but the sap in any one tube flows in only one direction at a time.
Must-know definitions
- Capillary adaptations
- Capillaries branch into huge networks with a very large total surface area; their narrow lumen (about 5–10 µm) brings every red blood cell close to the wall, which is one endothelial cell thick with pores between cells. Fenestrated capillaries also have pores through the cells, for especially rapid exchange.
- Artery
- Carries blood away from the heart at high, pulsing pressure. It has a thick wall relative to its narrow lumen: a thick tunica media of circular smooth muscle and elastic fibres, which stretch with each pulse and recoil between beats, and a collagen-rich tunica externa.
- Vein
- Carries blood back to the heart at low pressure with no pulse. It has a thin, flexible wall relative to its wide lumen, and pocket valves that prevent backflow when surrounding skeletal muscles squeeze the vein.
- Coronary occlusion
- Blockage of a coronary artery, usually when a blood clot forms on a fatty plaque (atheroma) that has already narrowed it. Cardiac muscle beyond the blockage receives no oxygen and dies: a myocardial infarction (heart attack).
- Correlation coefficient (r)
- A number from −1 to +1 that measures how closely two variables are linearly related: the sign gives the direction and the size gives the strength. Even a strong correlation does not prove that one variable causes the other.
- Transpiration pull
- Evaporation of water from the cell walls of leaf cells draws water out of the xylem by capillary action, putting the water column under tension (negative pressure). Cohesion between water molecules transmits the pull down the continuous column to the roots.
- Xylem vessel
- A continuous tube of dead cells joined end to end, with no cell contents and no (or incomplete) end walls, so flow is unimpeded. Lignified walls resist collapse under tension, and pits let water in and out.
- Tissue fluidAHL
- Fluid formed when blood pressure at the arteriole end of a capillary forces plasma out through the capillary wall (pressure filtration). It has much the same small solutes as plasma but very little protein. Most drains back into the capillary at the lower-pressure venule end.
- Double circulationAHL
- Blood passes through the heart twice per circuit: the right side pumps it to the lungs (pulmonary circulation, low pressure) and the left side pumps it to all other organs (systemic circulation, high pressure). Bony fish have a single circulation: heart → gills → body → heart.
- Root pressureAHL
- A positive pressure in root xylem caused by active transport of mineral ions into the xylem, so that water follows by osmosis. It moves sap upwards when transpiration is too slow, for example in very humid air or in spring before leaves open.
- Sieve tube and companion cellAHL
- Sieve tube elements are living cells with sieve plates, very little cytoplasm and no nucleus, so sap flows easily. Each is linked by plasmodesmata to a companion cell, whose nucleus and many mitochondria support active loading and unloading of sucrose.
Common exam mistakes
Loses marks Arteries carry oxygenated blood and veins carry deoxygenated blood.
Write instead Arteries carry blood away from the heart and veins carry blood towards it. The pulmonary artery carries deoxygenated blood and the pulmonary veins carry oxygenated blood.
Loses marks Capillary walls are thin because they are made of a thin membrane.
Write instead The capillary wall is a single layer of thin endothelial cells (each with two plasma membranes in the diffusion path) on a basement membrane. Say "one cell thick".
Loses marks The elastic fibres in artery walls contract to pump blood along.
Write instead Elastic fibres are not contractile. They are stretched by the surge of blood in systole and recoil passively in diastole, which keeps the pressure up between beats. Only the smooth muscle contracts, and it changes the width of the lumen.
Loses marks Measuring the pulse with your thumb, or multiplying a 15 second count by 15.
Write instead Use the index and middle fingertips (the thumb has its own pulse) on the radial or carotid artery, and multiply a 15 second count by 4 (or a 30 second count by 2) to get beats per minute.
Loses marks The strong correlation between saturated fat intake and coronary heart disease proves that saturated fat causes it.
Write instead A correlation coefficient shows only how strongly two variables vary together. Another factor linked to both (a confounding variable) could be the cause; establishing causation needs other evidence, such as controlled experiments and a plausible mechanism.
Loses marks Water is pushed up the xylem by the roots, which use ATP to pump it.
Write instead In a transpiring plant water is pulled up passively under tension, by evaporation from the leaves. No ATP is used to move water; the xylem vessels are dead. (Root pressure, AHL, uses active transport of ions, not of water.)
Loses marks In a dicot root, xylem and phloem form vascular bundles in a ring, as in the stem.
Write instead In the root the vascular tissue is central: xylem forms a star or cross shape with phloem between its arms. Vascular bundles in a ring are the arrangement in the stem.
Loses marks Drawing the heart with the left ventricle on the viewer's left.
Write instead Diagrams show the heart from the front, so the heart's left side (left atrium, thick-walled left ventricle, aorta) is on the viewer's right.
AHLLoses marks The atrioventricular valves close when the atria contract, and the semilunar valves open as soon as the ventricles start contracting.
Write instead Valves open and close because of pressure differences. AV valves close when ventricular pressure rises above atrial pressure; semilunar valves open only when ventricular pressure exceeds arterial pressure, so there is a short period with both closed.
AHLLoses marks Phloem sieve tubes are dead, like xylem vessels.
Write instead Sieve tube elements are alive: they keep a plasma membrane, which is needed for loading sucrose, even though they have lost their nucleus and most organelles. Companion cells keep them alive.
AHL
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Multiple-choice questions
Flashcards
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Give three adaptations of capillaries for exchange.
Branching networks and narrow diameter give a very large surface area; a wall one thin endothelial cell thick (short diffusion distance) with pores between cells; and, in fenestrated capillaries (e.g. kidney glomerulus, intestinal villi, endocrine glands), extra pores through the cells for very rapid exchange.
How do you tell an artery from a vein in a micrograph?
Compare wall thickness with lumen diameter. Artery - thick wall, narrow round lumen, thick tunica media, often a folded lining. Vein - thin wall, wide and often flattened lumen, thin tunica media.
How do the elastic and collagen fibres in artery walls help with high pressure?
Elastic fibres are stretched by each surge of blood in systole and recoil in diastole, pushing blood on and keeping pressure up between beats (about 80 mmHg). Collagen fibres are strong in tension and stop the artery bulging or bursting.
How do you measure heart rate from the pulse?
Place the index and middle fingertips on the radial pulse (wrist, thumb side) or carotid pulse (neck, beside the windpipe), count beats for 15 s and multiply by 4 (or 30 s × 2). Repeat and take a mean.
Two adaptations of veins for returning blood to the heart
Pocket valves close if blood starts to flow backwards; thin, flexible walls let contracting skeletal muscles squeeze the vein, pushing blood towards the heart.
Causes and consequences of coronary artery occlusion
Plaque (atheroma) builds up in the artery wall and narrows the lumen; a clot can form on a ruptured plaque and block it. Cardiac muscle beyond gets no oxygen and dies - a myocardial infarction (heart attack). Narrowing alone can cause angina.
What does a correlation coefficient tell you, and what can it not show?
Its sign shows whether the correlation is positive or negative and its size (0 to 1) how strong it is. Even a strong correlation does not prove that one variable causes the other, because of confounding variables.
Explain the transpiration pull in four steps.
Water evaporates from leaf cell walls and diffuses out of stomata; capillary action draws water through cell walls from the xylem; this creates tension (negative pressure) in the xylem; cohesion transmits the tension down the continuous water column to the roots.
Four adaptations of xylem vessels
No cell contents (dead, empty tubes); no or incomplete end walls (unimpeded flow); lignified walls (resist collapse under tension); pits (water enters and leaves).
Positions of xylem and phloem in a dicot stem vs root
Stem - vascular bundles in a ring near the outside, phloem outside and xylem inside each bundle, cambium between, pith in the centre. Root - central xylem shaped like a star or cross, with phloem between the arms, inside the endodermis.
How is tissue fluid formed and reabsorbed?
High blood pressure at the arteriole end forces plasma out through the capillary wall (pressure filtration), leaving cells and most proteins behind. At the venule end pressure is lower, and fluid drains back in, drawn by the plasma proteins retained in the blood.
Plasma vs tissue fluid
Tissue fluid has similar concentrations of water, ions, glucose and amino acids but very little protein and no red blood cells or platelets. Leaving the tissues it has less O₂ and nutrients and more CO₂ and urea.
How is excess tissue fluid returned to the blood?
It enters blind-ended lymph capillaries through gaps in their thin walls and flows as lymph through lymph ducts with valves, which empty into veins near the heart.
Single vs double circulation
Fish - heart → gills → body → heart (blood passes through the heart once per circuit; low pressure after the gills). Mammal - right side → lungs → left side → body → right side (twice per circuit; separate low-pressure pulmonary and high-pressure systemic circulations).
Why is the wall of the left ventricle thicker than that of the right ventricle?
The left ventricle must generate a much higher pressure to pump blood through the aorta to all the organs of the body. The right ventricle pumps only to the nearby lungs at lower pressure, which also protects the delicate alveolar capillaries.
Events on the left side of the heart in one cardiac cycle
SAN fires; atrial systole tops up the ventricle through the open bicuspid valve. Ventricular systole - bicuspid valve closes, then aortic valve opens and blood is ejected. Diastole - aortic valve closes, then bicuspid valve opens and the ventricle refills.
How is root pressure generated, and when is it needed?
Root cells actively transport mineral ions into the xylem; water follows by osmosis, creating a positive pressure that pushes sap up. It is needed when transpiration is too slow, e.g. in very humid air or in spring before deciduous plants have leaves.
How do companion cells and sieve tubes work together in translocation?
Companion cells (nucleus, many mitochondria) actively load sucrose, which passes through plasmodesmata into sieve tubes. Water follows by osmosis, raising pressure; sap flows through sieve plates (in cells with no nucleus and little cytoplasm) to sinks, where sucrose is unloaded.
Last reviewed 19 October 2026. Spotted a mistake? Report it.