Module 3: Exchange and transportTransport in animals (3.1.2)

Both surface area and volume are crucial factors in determining how substances move into, out of, and within organisms.

Surface area is the total external area of the organism that is in contact with the environment.

Volume is the total internal space within an organism.

As organism size increases, its surface area-to-volume ratio decreases.

Single–celled organisms have a high surface area-to-volume ratio, enabling efficient diffusion of nutrients, gases, and waste owing to short diffusion distances.

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Multicellular organisms can be large and complex, with specialised tissues and organ systems. Metabolic demands include taking in nutrients, exchanging gases, and excreting waste products.

Mass transport systems are needed to overcome the limitations of diffusion in moving substances (e.g., hormones, gases, nutrients, wastes) long distances between exchange sites and cells.

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Mass flow is the bulk movement of substances driven by a pressure difference. It allows rapid, directed transport over long distances in large organisms.

Diffusion occurs over short distances at exchange surfaces, where substances move down concentration gradients.

Mass transport systems link these exchange surfaces, enabling efficient distribution of gases, nutrients and wastes. They help maintain concentration gradients, ensuring effective diffusion and supporting normal cellular function.

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Circulatory systems provide a means of mass transport in multicellular organisms.

They consist of a:

  • transport medium
  • system of vessels
  • mechanism to generate pressure and drive movement.

Circulatory systems can be open or closed.

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In open circulatory systems, haemolymph is pumped into body cavities (haemocoel) rather than remaining within a closed network of vessels. This occurs in arthropods and many molluscs, where flow is low-pressure and less directed than in closed systems.

In insects, a tubular heart pumps haemolymph forward. It is released into the haemocoel, where it bathes tissues before returning to the heart via valves. Flow is maintained by peristaltic contractions of the body.

Oxygen delivery is largely independent, as tracheae supply oxygen directly to tissues.

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In an open circulatory system, the transport fluid (haemolymph) leaves the vessels and directly surrounds tissues.

In a closed circulatory system, the transport fluid (blood) remains within a continuous system of vessels. They work at higher pressure and are more efficient (faster and more controlled).

Closed circulatory systems can be single or double, depending on how many times blood passes through the heart in a circuit of the body.

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In single circulatory systems, such as in fish, blood passes once through the heart during a complete circuit of the body:

  • Deoxygenated blood is pumped to the gills for gas exchange.
  • Oxygenated blood then circulates through the body tissues, delivering oxygen and nutrients, and returns, deoxygenated to the heart.

It is less efficient than double circulatory systems as blood loses pressure after the gills.

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In a double circulatory system such as in mammals and birds, blood passes twice through the heart per circuit.

The right side of the heart pumps deoxygenated blood to the lungs (pulmonary circulation) while oxygenated blood returns to the left side of the heart and is pumped throughout the body (systemic circulation).

The heart pumps oxygenated blood at high pressure through a network of vessels to body tissues, ensuring large surface area for diffusion, short diffusion distances, steep diffusion gradients for fast oxygen and nutrient delivery.

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The composition of blood vessels varies by function. The presence and relative thickness of each layer differs between vessels.

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Arteries transport blood away from the heart rapidly and at high pressure.

Arteries have thick walls, muscle layers and elastic layers compared with veins:

  • Collagen walls restrict the maximum stretch of the vessel as blood is pumped under high pressure.
  • Smooth muscle controls blood flow through constricting and relaxing the vessel.
  • Elastic fibres maintain high blood pressure throughout the arteries ensuring blood can travel long distances, smoothly and quickly.
  • No valves (other than in the arteries at the point where they leave the heart).
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Arteries branch into narrower vessels called arterioles that link the arteries to capillaries.

  • Thicker muscle layer than in arteries to facilitate vasoconstriction and vasodilation.
  • The elastic layer is thinner than in arteries as the blood pressure is lower.
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Capillaries are a network of small vessels connecting arterioles to venules and surrounding tissues.

  • Their walls are a single layer of endothelial cells reducing diffusion distance between blood and tissue cells and increasing diffusion speed.
  • Small diameter slows blood flow, increasing the time available for diffusion and maintaining high pressure and a steep concentration gradient.
  • Gaps between endothelial cells allow cells and substances to pass out of the vessels and into the fluid surrounding tissues (e.g., white blood cells).
  • Highly branched network create larger surface area for metabolite exchange.
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Venules connect capillaries to veins.

They have a thin elastic layer, thin muscle layer and a large lumen owing to blood flowing at lower pressure after it has passed through the capillaries.

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Veins transport blood back to the heart, usually at low pressure.

  • Low blood pressure means there is only a need for a thin collagen layer. This also enables compression of vessels by surrounding muscles to generate pressure for circulation.
  • Thin muscle layer compared with arteries.
  • Thin elastic layer owing to low pressure within the vessel meaning stretch and recoil are not needed.
  • Valves are present throughout veins to ensure that blood flowing at low pressure moves only in one direction.
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Blood is composed of about 45 cells (red blood cells, white blood cells) and cell fragments (platelets) and 55 liquid plasma.

Plasma is about 95 water, making it a good solvent for transporting substances around the body.

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As blood flows through capillaries, some plasma leaks out through gaps in the capillary walls to surround body cells, forming tissue fluid.

Tissue fluid facilitates the exchange of substances including amino acids, ions, glucose, oxygen, carbon dioxide and other waste products between cells and the blood. Large plasma proteins are too big to pass through capillary walls and remain in the blood.

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Hydrostatic pressure is exerted by blood on the walls of the capillaries as a result of heart contractions.

At the arterial end of a capillary, hydrostatic pressure is high enough to push fluid out of the capillaries to become tissue fluid. Large proteins and cells remain in the blood. This is called ultrafiltration (filtration under pressure).

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Plasma proteins in the blood lower its water potential, causing water in the tissue fluid to move back into the blood by osmosis. This is called oncotic pressure.

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At the venous end of capillaries, hydrostatic pressure falls because tissue fluid moved out of the vessel, lowering the blood volume, and the pressure from heart contractions decreases with distance from the heart.

The reduced hydrostatic pressure relative to the oncotic pressure (unchanged) at the venous end of capillaries results in approximately 90 of tissue fluid re-entering the capillaries by osmosis having exchanged substances with tissue cells.

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The tissue fluid that does not return to capillaries (10) drains into lymph vessels and forms lymph fluid.

Lymph is similar to plasma and tissue fluid but contains less oxygen and nutrients (which have been exchanged with tissues) and more fatty acids absorbed from the small intestine.

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Lymph vessels are like blind–ended capillaries with one–way valves and form a network over the whole body, merging with larger vessels.

Fluid is moved around the vessels by the contraction of skeletal muscles.

The lymphatic system joins the bloodstream again via the subclavian veins.

Lymph nodes are distributed throughout the system. These contain lymphocytes and phagocytes that help fight infection.

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The mammalian heart is a muscular organ made of two pumps and is situated in the thoracic cavity. The heart is encased in the pericardium, a membrane that prevents the heart from overextending.

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The aorta distributes oxygenated blood from the left ventricle to the rest of the body.

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The vena cava are veins that carry deoxygenated blood from the head and the rest of the body back to the heart via the right atrium.

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In almost all cases, veins bring deoxygenated blood back to the heart and arteries take oxygenated blood away from the heart.

One exception is the vessels taking blood between the heart and lungs:

  • The pulmonary vein brings oxygenated blood from the lungs to the left atrium.
  • The pulmonary artery delivers deoxygenated blood from the right ventricle to the lungs.
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The coronary arteries are located on the surface of the heart and supply blood to the heart muscle.

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The mammalian heart consists of four chambers:

  • Blood enters the heart via the atria (upper two chambers) as it returns from the body (right atrium) and the lungs (left atrium).
  • The lower two chambers are the ventricles, which pump blood to the lungs (right ventricle) and to the rest of the body (left ventricle).
  • The septum is a muscular wall that divides the heart into the left and right sides. This separation prevents the mixing of oxygenated (left side) and deoxygenated blood (right side).
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Heart valves regulate blood flow and maintain pressure within the chambers. They ensure blood travels in the correct direction, avoiding backflow.

The pressure from blood flowing into the atria opens the atrioventricular valves. When the atria and ventricles are full, the atria contract, forcing blood down into the ventricles.

The contraction of the ventricles causes the atrioventricular valves to close, preventing backflow of blood into the atria. Blood is forced out of the ventricles through the semilunar valves in the aorta and pulmonary artery. These valves prevent blood from flowing back into the heart.

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Dissections are crucial for examining the internal structures of organs, allowing scientists to form theories about their functions and study anatomical relationships and comparative anatomy.

Hearts from sheep or pigs are often available from butchers and are similar in size to a human heart. They commonly have vessels and atria removed.

Ethical concerns include the treatment and sourcing of animals used for dissection, as well as adherence to cultural or religious beliefs.

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The cardiac cycle is the sequence of events that occurs within one complete heartbeat:

  • Atrial systole – the atria contract
  • Ventricular systole – the ventricles contract
  • Cardiac diastole – the heart relaxes
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In diastole, the heart is relaxed as it fills with blood.

As the atria fill with blood, the pressure exceeds that in the ventricles and the atrioventricular valves open so that blood fills the ventricles.

The pressure in the relaxed ventricles is lower than in the aorta and pulmonary artery, causing the semilunar valves to close.

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During atrial systole, the atria contract to force all the blood into the relaxed ventricles.

During ventricular systole, the thick ventricle walls contract and the pressure in the ventricles increases above that in the aorta and pulmonary artery. This closes the atrioventricular valves and blood is pumped to the lungs (right side) and the rest of the body (left side).

The pressure in the heart is low after ventricular systole but at a maximum in the arteries.

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Cardiac output is the volume of blood pumped per minute.

Where:
Stroke volume = the volume of blood pumped from the left ventricle in a single beat
Heart rate = the number of beats per minute

Units of cardiac output are volume per unit time (e.g. ).

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Cardiac muscle is myogenic, meaning that it contracts on its own without nervous stimulation. The heartbeat is initiated by electrical activity within the heart muscle itself.

Small changes in the electrical charge (polarity) of cardiac muscle cells spread as a wave of electrical excitation from cell to cell, causing them to contract in a coordinated sequence that produces the rhythmic heartbeat.

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The coordination of a heartbeat is a multistep process:

  1. The Sinoatrial node (SAN) initiates an electrical impulse, causing a wave of depolarisation over the atria (atrial systole).
  2. The atrioventricular node (AVN) receives the impulse and delays its spread to allow the ventricles to fill with blood.
  3. The impulse travels down the bundle of His and reaches the apex of the ventricles.
  4. Purkyne fibres conduct the impulse triggering ventricular contraction (ventricular systole) from the apex upwards, forcing blood into the pulmonary artery and aorta.
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An electrocardiogram (ECG) trace records the heart’s electrical activity.

Electrodes are placed on the skin to detect electrical signals generated by the heart.

ECGs help diagnose heart disorders by analysing the shape and timing of the waves.

Regular ECG monitoring helps track heart health and assess the effectiveness of treatments for cardiac problems.

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ECG’s can be used to diagnose normal and abnormal heart activity. A normal ECG shows a regular pattern of waves at a resting rate of 60–100

  • Tachycardia – resting heart rate above 100 The ECG shows normal wave patterns but they are closer together, with shorter intervals between beats.
  • Bradycardia – resting heart rate below 60 The ECG shows normal wave patterns but they are further apart, with longer intervals between beats.
  • Ectopic heartbeat (arrhythmia) – an extra or early beat caused by an electrical impulse originating somewhere other than the SAN. The ECG shows an additional or irregular wave disrupting the normal rhythm.
  • Atrial fibrillation – the atria contract rapidly and irregularly. The ECG shows no first wave and an irregular pattern subsequently.
  • Ventricular fibrillation – the ventricles contract rapidly and uncoordinatedly, so the heart cannot pump blood effectively. The ECG shows chaotic, irregular waves with no distinguishable structure. This is life-threatening and requires immediate defibrillation.
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Oxygen is primarily transported in the body by binding to haemoglobin, a protein found in red blood cells (erythrocytes).

Haemoglobin is a protein with quaternary structure composed of four polypeptide chains each containing a haem prosthetic group.

Each haem group contains an iron () ion that can bind one molecule of oxygen, allowing a single haemoglobin molecule to carry up to four oxygen molecules (eight oxygen atoms).

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When oxygen binds to haemoglobin, it forms oxyhaemoglobin. The reaction can be represented as:

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The binding of the first oxygen molecule induces a conformational change in the haemoglobin structure, increasing its affinity for oxygen and making it easier for subsequent oxygen molecules to bind.

The transition from a low-affinity to a high-affinity state is known as cooperative binding or positive cooperativity.

The reverse occurs when oxygen dissociates in tissues, shifting haemoglobin back to the low-affinity state, which promotes further oxygen release where it’s needed, e.g., where the partial pressure of oxygen is low.

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Carbon dioxide is a waste product of cellular respiration that diffuses from tissues into the blood and is transported in three main ways:

  • A small proportion (~5-10) of dissolves directly in blood plasma and is transported in solution.
  • can bind to haemoglobin, forming carbaminohaemoglobin.
  • The majority of is transported in the form of hydrogen carbonate ions, formed in red blood cells.
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The majority of is transported as hydrogencarbonate ions.

diffuses into the blood and then into red blood cells (erythrocytes). Inside red blood cells, carbonic anhydrase catalyses the reaction of with water to form carbonic acid ():

Carbonic acid then dissociates into hydrogen ions () and hydrogencarbonate ions ():

The ions bind to haemoglobin, forming haemoglobinic acid (HHb). This buffers the blood and facilitates oxygen release (the Bohr effect).

The ions diffuse out of red blood cells into the plasma via a transport protein. To maintain the electrical balance across the red blood cell membrane, chloride ions () diffuse into red blood cells from the plasma – this is the chloride shift.

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The oxygen dissociation curve shows the relationship between:

  • Partial pressure of oxygen (), which refers to the pressure exerted by oxygen in a mixture of gases.
  • Haemoglobin saturation, which refers to the percentage of haemoglobin’s four oxygen-binding sites that are occupied. When all four sites are bound to oxygen, haemoglobin is fully saturated.

The shape of the curve reflects haemoglobin’s changing affinity for oxygen: as each oxygen molecule binds, it becomes easier for the next to bind, producing the characteristic S-shaped sigmoid curve.

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Initial slow binding: At low the first oxygen molecule binds slowly because haemoglobin is predominantly in a low–affinity state, resulting in a shallow gradient at the bottom left of the curve.

Cooperative binding: Once the first oxygen binds, haemoglobin undergoes a conformational change, increasing its affinity for subsequent oxygen molecules. Saturation increases rapidly relative to causing a steep gradient in the curve.

Approaching saturation: As haemoglobin becomes nearly fully saturated, fewer binding sites remain available, so additional oxygen molecules bind more slowly, resulting in the curve levelling off at the top right.

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The Bohr effect describes how haemoglobin’s oxygen affinity decreases in response to high levels and low

In respiring tissues, diffuses into red blood cells, where the enzyme carbonic anhydrase catalyses its reaction with water to form carbonic acid (). This rapidly dissociates into and hydrogencarbonate ions (). The increased concentration lowers the causing haemoglobin to change shape and reduce its affinity for oxygen.

The Bohr effect shifts the oxygen dissociation curve to the right, meaning haemoglobin releases more oxygen at a given

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Foetal haemoglobin (HbF) is produced in the red blood cells of a foetus and binds oxygen from maternal blood across the placenta. HbF has a higher oxygen affinity than adult haemoglobin (HbA).

The HbF oxygen dissociation curve is shifted to the left of the HbA curve, indicating higher oxygen saturation at the same

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