Quiet breathing chiefly uses active inspiration and passive expiration; deeper or more forceful breathing recruits additional muscles and changes their coordination. The vertical, anteroposterior and transverse dimensions of the thorax can all change, and different patterns need not represent different mechanisms of gas exchange.
The respiratory diaphragm is the principal inspiratory muscle. Its contraction lowers and flattens its domes, increases the thoracic vertical dimension and displaces abdominal contents so the relaxed abdominal wall moves outwards. Its lower-rib attachments also contribute to expansion of the costal margin. The phrenic nerves, chiefly C3–C5, provide its motor supply. The external intercostal muscles elevate and stabilize the ribs, while the interchondral or parasternal portions of the internal intercostal muscles also assist inspiration. It would be wrong to give every internal intercostal fibre an exclusively expiratory function.
Elevation of upper ribs and the sternum increases the anteroposterior dimension, the pump-handle component. Outward movement of the lateral portions of lower ribs increases transverse diameter, the bucket-handle component. These are components of oblique rib rotation rather than two mechanically isolated systems. Abdominal or diaphragmatic breathing makes descent of the respiratory diaphragm particularly conspicuous; costal breathing makes chest-wall expansion more conspicuous. Both contribute in ordinary respiration, with their proportions changing with posture, mechanical loading and individual anatomy.
As the thorax expands, pleural pressure becomes more negative and transpulmonary pressure increases. The lungs expand and alveolar pressure briefly falls below atmospheric pressure, admitting air. Relaxation of inspiratory muscles lets elastic recoil reverse the flow in quiet expiration. The abdominal wall and respiratory diaphragm return towards their resting arrangement without requiring active abdominal compression for each quiet breath.[ref]
22.3 The Process of Breathing. Mechanisms of Breathing; Pulmonary Ventilation: pressure changes and inspiratory versus expiratory activity
With deeper inspiration, the scalene muscles help stabilize or elevate the upper ribs, and the sternocleidomastoid muscle can elevate the sternum when the head is fixed. With the upper limbs or shoulder girdle fixed, muscles such as pectoralis minor and serratus anterior can assist expansion in greater effort. Recruitment is graded; an anatomical list of possible accessory muscles does not mean all are active in every ordinary breath.
Active expiration uses the abdominal muscles, including rectus, oblique and transversus components, to compress the abdominal contents and drive the respiratory diaphragm upwards. The interosseous portions of the internal intercostal muscles help depress the ribs. This is important in vigorous exercise and expulsive actions. At rest, elasticity supplies much of the expiratory work; during high demand, active muscle contraction adds to it.
The observable pattern also changes with the physiological task. Exercise increases breath depth and usually frequency. A sigh is an enlarged inspiration followed by expiration; yawning includes a prolonged deep inspiratory movement. Coughing coordinates an inspiration, closure of the glottis, pressure-generating expiratory effort and sudden glottic opening. Speech imposes a controlled prolonged expiration with laryngeal regulation. Swallowing briefly interrupts ordinary airflow. Thus respiratory observation should include phase, depth, frequency and coordination, not merely count which ribs move. Fixed claims that men must breathe abdominally and women must breathe costally overstate a variable tendency. Pregnancy or a change of posture can alter mechanical contributions without replacing the basic pressure mechanism.
The defensible core of William Odling's statement is a transfer of energy: sunlight helps build reduced food material, and its subsequent oxidation can supply animal heat and work. Here solar force means solar energy, and eremacausis means slow oxidation, rather than a special vital agent. The historical wording, from his 1866 lectures, must be qualified where it identifies the fuel with the substance of the working muscle or contrasts plant and animal metabolism too absolutely.[ref]
Lectures on Animal Chemistry by William Odling. Lecture IV, discussion of solar energy and animal oxidation; see also the concluding recapitulation
In oxygenic photosynthesis, light absorbed by chlorophyll supports conversion of carbon dioxide and water into reduced organic material, with release of oxygen. An overall account, written for one glucose unit, isThis summarizes several stages, rather than describing a single elementary chemical reaction. The evolved oxygen comes from water. The product can furnish starch, other carbohydrates and precursors for fat and protein; the latter additionally requires a source of nitrogen. A human obtains these materials directly from plants or through animals that have eaten plants. Thus a grain meal and the flesh of a grain-fed animal can trace much of their chemical energy to the same original sunlight.[ref]
8.1 Overview of Photosynthesis. Main Structures and Summary of Photosynthesis; photosynthetic products and their passage through food chains
The reverse overall balance for complete aerobic utilization of glucose isSimilarly a representative fatty acid, palmitic acid, has the complete oxidation balanceBoth equations conserve each chemical element. The relevant energy difference belongs to the complete set of reactants and products, including oxygen; merely breaking a bond does not supply net energy. In modern terms, controlled cellular respiration couples part of the available chemical free energy to ATP formation. ATP hydrolysis can then support muscle contraction, active transport and biosynthesis; some energy is transferred to heat. The process occurs through enzyme-mediated steps, without a flame, although complete combustion in a calorimeter can help measure the corresponding chemical energy change.
Glands and muscles continually replace constituents, and their proteins can be degraded. Amino acids can contribute carbon skeletons to energy metabolism, while their nitrogen is disposed of largely as urea. But ordinary muscle contraction does not require proportional destruction and rebuilding of the contractile protein at every stroke. Carbohydrates and fats are major fuels for muscle, and a gland can use those fuels to support secretion. The historical picture of burning the organ's own substance is therefore an inadequate general explanation.[ref]
This correction was already an experimental issue before the exam. Edward Frankland's 1866 discussion compared the mechanical work of climbing with the energy that could be assigned to protein destruction estimated from urinary nitrogen. The latter was insufficient to account for all the work. His combustion measurements also distinguished the heat from burning a sample completely from the smaller amount available when nitrogenous material leaves the animal as still-combustible urea. Measuring external work, urinary products and calorific values gives the hypothesis a quantitative test; it is more informative than simply pointing out that an active animal gets warm.[ref]
On the Source of Muscular Power by Edward Frankland. pp. 394–395, criticism of tissue-consumption doctrine; pp. 401–406, calorific measurements and comparison with climbing work, 1866
Finally, plants themselves carry out cellular respiration, and not every ecosystem obtains its input from sunlight: chemolithotrophy is an alternative. The plant-animal contrast is useful for following energy through an ordinary food chain, but it is not an exhaustive division of biochemical capabilities. Conservation of energy also requires the bookkeeping formfor an idealized interval in which is the metabolizable food input, with energy lost in excreta already subtracted and other material-energy exchanges neglected. Here is external work, is net outward heat transfer, and includes changes in stored chemical and other energy. The living body transforms energy; it does not create it.
The heart supplies the principal continuing mechanical input to blood circulation; arterial recoil, respiratory movements and the action of skeletal muscles redistribute and supplement that input. They have different sources of energy and should not all be called independent propulsive powers. The pressure gradient remaining after passage through the capillaries is still a consequence of the upstream pump.
Arterial elasticity. During ventricular systole, ejected blood expands the aorta and large arteries, storing elastic potential energy in their walls. During diastole, recoil continues to drive blood into the peripheral vessels while closure of the aortic valve limits return towards the heart. This reduces the intermittency of flow downstream. The recoil is a temporary store of cardiac work, rather than a fresh source of energy. A vessel that is too rigid provides less of this smoothing; a compliant vessel with an open backward outlet would not provide the same useful result.
The skeletal-muscle pump. Contracting skeletal muscles compress adjacent veins. Venous valves make the displacement preferentially towards the heart; when compression ceases, refilling permits another stroke. Walking therefore aids return from dependent limbs. The source of the added work is muscle contraction, and the valves guide it without producing energy. Merely possessing valves cannot make a stationary column circulate.
The respiratory pump. During inspiration, falling intrathoracic pressure and displacement of the abdominal contents favour a pressure gradient from abdominal and peripheral veins towards the thoracic veins and right side of the heart. The respiratory diaphragm thus participates indirectly in venous return as well as pulmonary ventilation. The exact flow varies with posture, vessel collapse and the phase of the respiratory cycle; positive-pressure inflation does not reproduce the same mechanical circumstances as spontaneous inspiration.[ref]
Venous tone and cardiac filling. Sympathetic contraction of venous smooth muscle can reduce venous storage capacity and transfer blood towards the central circulation. This alters the volume available for filling rather than requiring a wave of venous peristalsis. Cardiac relaxation, recoil and movement of the atrioventricular plane also affect filling pressure. Some traditionally described cardiac suction is therefore a feature of the cardiac cycle and mechanical coupling, not a separate organ pulling blood through every vessel.[ref]
20.4 Homeostatic Regulation of the Vascular System. Neural Regulation of Blood Flow; sympathetic effects on vessels and venous return
Gravity and supposed capillary attraction. Gravity assists a downward column but opposes an upward one. In a complete circuit returning to its original height it supplies no net gravitational work; its important practical effects include hydrostatic pressure differences and dependent venous pooling. Ordinary capillary action in an open narrow tube, driven by a liquid meniscus, is not a general pump in a fluid-filled capillary network. Nor does a vague attraction of blood by tissues replace a measurable pressure gradient. Local vasodilation can increase perfusion by lowering vascular resistance, but that change does not manufacture the necessary pressure source.
For a simple mean-flow description, with the volume flow rate, the driving pressure gradient and the effective vascular resistance,This relation distinguishes a source of driving pressure from a change of resistance. In a lumped compliant arterial reservoir with vascular compliance and no inflow between beats, volume balance giveswhere is the downstream pressure. The exponential decay follows from stored volume leaving through the resistance. It explains continued flow during diastole without claiming that the idealization describes every reflected arterial wave.
A physical demonstration can compare a pulsatile pump feeding a narrow outlet through rigid tubing with the same pump feeding it through an elastic reservoir. Collecting the outflow over the pumping and resting phases reveals the reservoir's smoothing action; a side-arm manometer records its falling pressure. Separately compressing a valved flexible tube shows the role of a peripheral muscle pump. Keeping pump stroke and outlet resistance comparable prevents the apparatus from conflating a smoother flow with creation of extra energy.