1 Cellular Physiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

2 Autonomic Nervous System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45

3 Neurophysiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65

4 Cardiovascular Physiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113

5 Respiratory Physiology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185

6 Renal Physiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239

7 Acid-Base Physiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303

8 Gastrointestinal Physiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 329

9 Endocrine Physiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383

10 Reproductive Physiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 447

Appendix I Common Abbreviations and Symbols . . . . . . . . . . . . . . . . . . . . 469

Appendix II Normal Values and Constants. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .471

Challenge Yourself Answers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 473

Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .477

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Physiology - 9781455708475 | US Elsevier Health Bookshop

Physiology is the branch of biology that deals with the functions and processes of living organisms, both animals and plants. It's biology in motion.

Physiology includes everything from how a single cell functions, to what makes your nerve receptors work, how your pancreas releases insulin, and what happens to your muscles when you exercise. Technology has made for great leaps in the science of physiology. The electron microscope, for instance, allows you to see down to even the molecules of the cell, and radioactive isotopes provide the means to track the movement of substances within an organism.

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physiology - Dictionary Definition : Vocabulary.com

This week marks the start of the 2017 Tour de France. This year the race covers 2,200 miles in 21 days of racing, comprised of team and individual time trials as well as stages through cities, countryside and mountains of France.

The Tour de France is interesting to me because it provides an excellent opportunity for a short lesson in sports physiology.

All of the riders in the Tour are exceptionally fit since their bodies have adapted to years of dedicated, intense training. Endurance sports like cycling are dependent on the delivery of oxygenated blood to the muscle to produce ATP, the energy needed to sustain exercise.

The riders have large, strong hearts, resulting in the ejection of more blood to the muscle. Within the muscle there is an increase in the number of capillaries, the small blood vessels that deliver blood to the muscle, and mitochondria, the part of the cell that produces most of the ATP. Together, these adaptations allow the muscle to produce more ATP without fatigue, allowing the athlete to exercise at a higher intensity for a longer time.

But training isnt the only reason these athletes can sustain such intense exercise for so long. Proper nutrition, especially what the athletes eat and drink before, during and after each stage, also plays an important role.

Intense endurance exercise like cycling relies on carbohydrates, in particular, muscle glycogen, as a fuel. Muscle glycogen is a storage form of glucose, sugar that the muscle converts into energy. During prolonged exercise that lasts several hours, muscle glycogen levels can be severely depleted.

Eating carbohydrates before exercise can boost muscle glycogen levels, so cyclists eat carbohydrate-rich foods for breakfast before each stage. They also consume carbohydrates in the form of sports drinks (think Gatorade) and energy bars prior to starting.

In fact, they start replenishing their muscle glycogen immediately after finishing the previous days ride. This usually begins with a recovery beverage, which may contain some protein for more rapid muscle glycogen synthesis, and extends through carbohydrate-rich meals and snacks that afternoon and evening.

During exercise it is crucial to maintain adequate blood glucose levels, which tend to drop since the muscle is using so much as a fuel. Failure to replenish blood glucose results in what cyclists call hitting the wall or bonking, which is like your car running out of gas. To prevent this, glucose must be replenished, typically with sports drinks, energy bars or a sugary mixture called goo.

Prolonged, intense exercise, especially in the heat, results in a high sweat rate, which can lead to dehydration. Sweat loss of several liters per hour is not uncommon during cycling, so fluid intake is essential. This means that cyclists spend a lot of time drinking water while they ride. Sports drinks are also commonly used since they contain carbohydrates and electrolytes in addition to water.

Endurance events like cycling, especially multi-stage events like the Tour de France, highlight important concepts of sports physiology. Even though you may never compete at that level, understanding how training can improve your endurance is relevant if you cycle or run, walk or swim for exercise.

Knowing how proper nutrition before, during and after exercise can improve performance can help you make better decision about what to eat. Hopefully, it also gives you a greater appreciation for the science that goes into a performance like the Tour de France.

Brian Parr, Ph.D., is an associate professor in the Department of Exercise and Sports Science at USC Aiken where he teaches courses in exercise physiology, nutrition and health behavior. You can learn more about this and other health and fitness topics at http://drparrsays.com or on Twitter @drparrsays.

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HEALTH AND FITNESS: Sport physiology in the Tour de France - Aiken Standard

Evacuation of Dr. Ronald S. Shemenski from the British Antarctic Survey Rothera Research Station, April 26, 2001. The airlift operation was the riskiest rescue effort ever by a small plane to the South Pole, as the weather makes any flights to the South Pole extremely hazardous from late Februray until November. (AP Photo/British Antarctic Survey) Photograph: AP

The International Geophysical Year was a global survey, but it had a particular impact on Antarctica, as it led to the creation and signing of the Antarctic Treaty, reserving the continent for peaceful purposes only and ensuring Freedom of scientific investigation. While most of the work done was as the name suggests in the physical and geographical sciences, one almost unknown part of the research involved an international team of physiologists and doctors who headed out to Antarctica to study the human body in an extreme environment.

INPHEXAN, the INternational PHysiological EXpedition to ANtarctica involved six researchers from three countries: Nello Pace, William Siri and Charles Meyers from the USA; Gerhard Hildebrand, a recent German immigrant to the USA (and ex-First Alpine Battalion member); and James Adams and Lewis Griffith Evans Cresswell Griff Pugh from the UK. Initiated by Pace and Siri, who shared leisure interests in high altitude climbing as well as research interests in stress and physiology, the initial plan was a study of hormonal responses to the stress of the Antarctic environment the cold, dark, and isolation. Charles Meyer, a dentist and bacteriologist at the Naval Biological Laboratory in Berkeley went along to conduct studies of infectious diseases. The UK team had intended to study changes in metabolism, and the possibility that people are able to acclimatise to intense cold, and agreed to join with the Americans to make an international research team.

One metabolic study involved a brand new piece of technology: the IMP, or Integrating Motor Pneumotachograph, invented by Heinz Wolff at the National Institute for Medical Research (before he started his TV career on The Great Egg Race and other shows). A variation on the Douglas Bag, the IMP measures the volume of air used by the human subject, and takes samples of their breath for analysis. The UK team measured the metabolism of several scientists, with the help of Allan Rogers, the Medical Officer on the Commonwealth Trans-Antarctic Expedition (TAE) of 1955-8.

Strapped into the IMP for a week, the geologist Geoff Pratt wrote up a report On being IMPed where he complained about feeling suffocated in the mask, that it got in the way when he was trying to work, and that it stopped him communicating effectively with his colleagues. He also blamed the suit, the tight mask, and the difficulty he had sleeping with it on for a string of accidents and mistakes in his lab

in the course of the week I have done a remarkable number of stupid things.

You can see the IMP in action in a short British Pathe Film (the IMP appears at 1.20).

While the team got relatively few publications out of this work it did enable researchers to improve and adapt the equipment to make it easier on human subjects. Pughs work with the copper-wire body suit (in the video above) and other measurements became papers on the effects of solar radiation on temperatures in the Antarctic, and he also published in Nature on the blood of Weddell seals, and on the dangers of Carbon Monoxide poisoning in explorers huts.

Allan Rogers other major investigation was what appeared to be a very simple study of acclimatization to cold: he gave all the members of the TAE cards to fill out, every day, to record their clothing, their sleep patterns, their activities, any illnesses, and any other information they thought relevant. After the 15 month expedition was over, he intended to discover if men who had spent a long time in Antarctica wore fewer clothes in other words, were more acclimatized to cold than new arrivals on the continent, using the data from around a dozen members of the expedition.

Unfortunately the task turned out to be anything but simple: at least three academic statisticians tried and failed to analyse the huge amount of data and correlate it with weather patterns, working patterns, and sickness records. Ten years after the IGY Rogers finally got the money to hire someone who could do the job: recent mathematics graduate Mrs RJ Sutherland, who designed a computer programme to deal with the pile of report cards and all their information. Finally, in 1971, Sutherland and Rogers published their report which showed a negative finding: the men of the TAE felt the cold just as much at the end of their trek as at the beginning.

The vision of women slowly crunching numbers in a computer room in Bristol, or men patiently filling in sleep cards or trying to get on with their jobs while wearing an IMP, might not be particularly exciting or glamorous, this was hard, boring, awkward work that mattered. The IGY provided crucial data that helped us understand how the planet worked and particularly gave us a baseline for understanding climate change and none of that would have been possible without physiologists and other biomedical scientists designing safe rations, comfortable snow goggles, and warm gloves.

A collection of Allan Rogers possessions including clothing, his IMP equipment, and his medical kit are in the collections held by the Scott Polar Research Institute. The Year That Made Antarctica: People, Politics and the International Geophysical Year is on display at the Institutes Polar Museum, University of Cambridge, until 9 September 2017.

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Blood, Sweat and Ice? During the 60th anniversary of the IGY lets ... - The Guardian

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John Knight is senior lecturer in biomedical science; Chris Wigham is senior lecturer in interprofessional studies; Yamni Nigam is associate professor in biomedical science; all at the College of Human Health and Sciences, Swansea University.

The special senses sight, hearing, smell, touch and balance allow us to perceive the world and communicate. Like all body systems, they undergo age-related changes that negatively affect their function. Physiological changes to the eyes and ears mean older people gradually see, hear and balance less well. The changes also increase the risk of conditions such as cataracts, age-related macular degeneration, and conductive and sensory hearing loss. This sixth article in our series on the effects of age on the body describes what happens to the eyes and ears.

Knight J et al (2017) Anatomy and physiology of ageing 6: the eyes and ears. Nursing Times [online]; 113: 7, 39-42.

The special sense organs the eyes, ears, nose, tongue detect information coming from the environment, such as light, sound, smells and tastes, which is then relayed to the brain where it is processed into meaningful sensations. Diminished acuity of the special senses reduces our ability to perceive the world and communicate. All the senses go through an age-related decline, but the most dramatic changes are seen in the eyes and ears. This sixth article in this series on the effects of ageing on the different body systems examines the age-related changes in the eyes and ears.

Vision is affected by the ageing of the internal and external structures of the eye. Its decline is gradual and linear, and detectable changes begin in the third decade of life. The main changes are outlined in Fig 1.

Anatomy and physiology fig 1

Anatomy and physiology fig 1

The retro-orbital fat, which protects and cushions the eye, atrophies with age, causing the eyeball to recede into its socket (enophthalmos). As a result, eyelid tissues become lax and the levator muscles in the eyelids weaken, causing the eyelids to droop (ptosis). Drooping eyelids can gradually obstruct the upper field of vision.

Sinking eyeballs and drooping eyelids often lead to the conjunctiva lining the eyelids (tarsal conjunctiva) failing to sufficiently lubricate the front of eye (cornea). This can result in an air space developing between the lid and the cornea, particularly at night, potentially leading to epithelial breakdown (Sobel and Tienor, 2013; Liang et al, 2011).

In some people, the weakening of the muscles supporting the eyelids and loosening of the eyelids result in the eyelashes turning inwards (entropion) and irritating the corneas surface, which could lead to ulceration. A significant weakening of the muscles supporting the lower eyelids can result in them flopping away from the eyeball (ectropion), which can then dry out and become irritated. Symptoms of entropion and ectropion can be relieved by anti-inflammatory eye drops or artificial tears, while surgery to tighten the skin and muscles of the eyelid can provide permanent relief (Garrity, 2016).

With age, the lacrimal glands produce fewer tears, while the composition of tears changes, and the wetting efficiency and stability of the tear film is reduced. This leads to dry eye syndrome in up to 14% of the over-65s. Irritation, grittiness and pain can ensue and affect many activities, such as reading or watching television. Persistent irritation can make the cornea less sensitive, which in turn can diminish the ability to detect injury or infection. Many people find the use of artificial tears effective (NHS Choices, Dry Eye Workshop, 2007).

Throughout life, equatorial lens cells divide and new cell layers are added to the outside of the lens. Since the lens cannot grow in size (if it did it would soon outgrow the eyeball), its cells must be compressed, which results in the lens becoming increasingly dense and inflexible. The lens, therefore, becomes progressively less able to change shape to be able to focus light on the retina.

Presbyopia is the age-related reduction in the ability to see near objects. It typically presents as an inability to read text positioned close to the eye and generally develops in the 40s and 50s. Presbyopia results from a reduction in the ability of the intraocular lens to change shape. The distance from the eye at which print can be read (near point) increases from about 10cm at the age of 20 to over 100cm by the age of 70. Most people manage presbyopia by using correcting reading glasses (Boyd, 2016).

New lens cells continue to be produced throughout life, so the lens continues to increase in density. This can cause particular light frequencies to be absorbed and the lens to take on a yellowish hue, affecting contrast sensitivity and the accurate perception of colours.

When the concentration of proteins in the lens becomes very high, precipitation occurs. This is seen as a cataract. Thescattering of the light causes a glare or a halo effect when looking at bright lights. As the densest area of the lens is the centre, this is where age-related cataracts are most commonly seen.

Cataracts can make it difficult to see in certain circumstances for example, when driving at night. They can also interfere with the ability of certain wavelengths of light to enter the eye, thereby reducing colour perception: people with cataracts may wear garish clothing due to their compromised colour vision.

Individuals with poorly controlled diabetes are at much greater risk of developing cataracts because increased blood glucose encourages the build-up of damaging levels of sorbitol in the lens (Knight et al, 2017).

Cataracts are managed by removing the lenss contents from the capsular bag and placing a small intraocular lens inside the capsule to provide refractive power. The power of the intraocular lens can be chosen to suit the patients wishes and lifestyle (Truscott, 2003).

One role of the pupils is to regulate the amount of light entering the eye. With age, their diameter decreases, reducing the admittance of light. Age also has a negative effect on the pupils ability to adapt to changes in light intensity for example, when going from light to dark. Adapting to the dark requires the photosensitive cells of the retina to regenerate the photopigment rhodopsin; this is considerably delayed with age, which contributes to night-vision problems.

These changes increase older peoples risk of falls and other accidents, for example, when leaving a brightly lit bathroom to walk up or down a flight of poorly lit stairs (Rukmini et al, 2017; Turner and Mainster, 2008; Bitsios et al, 1996).

The eye consists of two hollow chambers separated by the lens. The anterior chamber is filled with a watery fluid (aqueous humor) and the posterior chamber with a jelly-like material (vitreous humor). The composition of the vitreous humor can change from a gel to liquid with age and, in some people, it shrinks, collapses and separates from the retina. This posterior vitreous detachment often manifests as discrete opacities (floaters) or sheering patterns in the field of vision (Bishop et al, 2004).

With age, cone photoreceptor cells in the fovea, which provide high-quality colour vision, begin to die, eventually resulting in age-related macular degeneration (ARMD). This is thought to be caused by changes to the cells of the retinal pigment epithelium (RPE), which lies next to, and maintains, the photoreceptor cells.

There are two types of ARMD: dry (90% of cases) and wet (10%). Dry or atrophic ARMD is characterised by a gradual bilateral loss of vision as the RPE degenerates. Wet or exudative ARMD is caused by the growth of new blood vessels in the space between photoreceptors and RPE (subretinal space) and the leakage of serous fluid from these new vessels. Wet AMRD has a more rapid onset and causes more severe loss of vision.

In ARMD, pale yellow-white elevated spots called drusen appear on the retinal surface, distorting vision and reducing visual acuity. Their appearance steadily increases after the age of 60 years. ARMD accounts for half of all visual impairments among people aged 75 and over (AMD.org; National Eye Research Centre; Forrester et al, 2001).

The ear is the organ of hearing but also plays the major role in our sense of balance. Problems with hearing are the most common sensory disorder associated with ageing. At age 61-70 years, around a third of people develop problems understanding speech if there is ambient background noise, and in those aged 85 years and over this rises to around 80% (Sogebi, 2015). Age-related changes to the ear are shown in Fig 2.

Anatomy and physiology fig 2

Anatomy and physiology fig 2

The auricle (pinna) collects sound waves and directs them through the ear canal (auditory meatus) to the eardrum. With age, the pinna often becomes larger and features more external hair on the tragus and lower helix; these changes are more often seen in men. The pinna becomes increasingly dry and scaly in both sexes.

The auditory meatus produces earwax (cerumen), which moistens the ear canal and is mildly antiseptic, helping to keep the ear free from infection. Unless compressed and pushed inwards by implements such as cotton buds, cerumen gradually works its way out (the ears are often described as self-cleaning).

With age, the ceruminous glands become less active and produce less earwax, which can lead to the auditory meatus becoming increasingly dry and prone to infection. The cartilaginous components that form the walls of the auditory meatus can lose elasticity, degrade and sometimes collapse, which increases the likelihood of ear canal collapse (Howarth and Shone, 2006). A drier environment and ear canal collapse both increase the likelihood of cerumen accumulation and obstruction, commonly resulting in conductive hearing loss. Older people may need to use earwax softeners before having excess wax removed by micro-suction at audiology clinics or by syringing at GP surgeries.

The middle ear consists of the ear drum (tympanic membrane) and a hollow, air-filled chamber spanned by three tiny bones (auditory ossicles):

The tympanic membrane vibrates in harmony with the sound waves collected by the outer ear, and these vibrations are transmitted and amplified across the middle ear by the three auditory ossicles. With age, the tympanic membrane becomes less vascular and begins to thin and stiffen (Liu and Chen, 2000; Weinstein, 2000). In older people, the tiny synovial joints between the three auditory ossicles are often stiff and calcified, leading to less efficient conduction and amplification of sound waves.

The air-filled chamber of the middle ear is connected to the back of the pharynx by the auditory or Eustachian tube: this ensures the pressure is kept relatively equal on both sides of the eardrum to prevent pressure building up and damaging the tympanic membrane. The musculature lining the auditory tube often undergoes age-related atrophy, which may interfere with the tubes opening during swallowing, thereby increasing the risk of pressure differences between the two sides of the eardrum.

The inner ear consists primarily of the:

The cochlear is a fluid-filled, spiral-shaped organ that receives sound waves directly from the stirrup. Sound waves travel rapidly through the fluid of the cochlear and are detected by special sensory receptor cells called hair cells. These relay auditory signals to the cochlear nerve, which delivers them to the auditory cortex of the brain, where they are perceived as sound. Our sense of hearing is most acute at the age of 10 years and gradually declines thereafter.

Almost everyone experiences a deterioration in hearing as they age, and currently there is no way of preventing or reversing these age-related changes. Presbycusis is the sum of all conditions that lead to decreased hearing sensitivity with age; it can be accelerated by exposure to loud noise, conditions that impair cardiovascular function and nerve damage (Parham et al, 2011). Presbycusis is usually associated with a progressive degeneration of the hair cells and neurones in the cochlea.

It has been suggested that a lifetime exposure to loud noises cumulatively damages hearing. Indeed, some people living in isolated, non-industrial communities in Africa and India have little age-related hearing loss. Inheritance of certain genes, increased exposure to free radicals and toxins, and decreasing blood supply to the inner ear (Danner and Harris, 2003) contribute to presbycusis and the rate at which it develops. A slowing in the brains processing of auditory information is another contributing factor.

Presbycusis is particularly associated with a declining ability to hear high frequencies, which are important for interpreting speech. As a result, older people find it increasingly difficult to follow and join in conversations, especially when competing background sounds (for example, from television or music) are present. This can restrict interactions and contribute to loneliness and social isolation (Parham et al, 2011).

Tinnitus is the hearing of a noise often a ringing, buzzing, humming or whooshing in the absence of any external sound; it is occasionally reported as having a musical quality. The condition has a variety of causes, including:

However, the major cause of tinnitus might be the lack of sensory input reaching the auditory cortex of the brain. Tinnitus has been compared to phantom sensations perceived in a non-existent limb after amputation: in some people, the sounds associated with tinnitus persist even after the cochlear nerve has been severed (Danner and Harris, 2003). As ageing is associated with a loss of sensory hair cells, the resultant reduction in sensory input to the brain may explain why prebycusis and tinnitus often coexist.

There is mounting evidence that exposure to loud sounds throughout life can both accelerate age-related hearing loss and increase the risk of tinnitus. It is a concern to audiologists that growing numbers of young people attend loud concerts and listen to loud music through headphones for long periods this is likely to accelerate their hearing loss and lead to hearing problems and deafness much earlier in life (Kujawa and Liberman, 2006).

The ability to balance the body at rest (static balance) and when moving (dynamic balance) relies on a complex interplay between different sensory systems including sight, touch and the vestibular system of the inner ear. To trigger the intricate motor coordination of skeletal muscles required to maintain balance, various regions of the brain need to quickly process a large and continuous input from these sensory systems (Horak, 2006).

The vestibular system of the inner ear consists of a labyrinth containing semicircular canals and their hair cells, and the otolith organs (utricle and saccule). All are key in maintaining balance. With age, the vestibular apparatus progressively loses hair cells some people aged 70 years or over experience up to 40% reduction in hair cells in the semi-circular canals (Rauch et al, 2001).

Other notable changes are the progressive fragmentation and degeneration of the otoliths (tiny stones made of calcium carbonate), particularly in the saccule. The number of vestibular nerve cells also diminishes from around the age of 60 years. These changes mean that, with age, our sense of balance becomes impaired and we may experience dizziness. Poor balance and dizziness, together with frailty and reduced reaction times, contribute to the risk of falls a major concern in older people. Each year an estimated 20-40% of those aged 65 and over fall at home (Shupert and Horak, 2017).

While little can be done to avoid the effects of ageing on sight and hearing, it is vital to encourage older people to have regular eye and hearing tests (Box 1). This means appropriate glasses and/or hearing aids can be dispensed, and common age-related pathologies such as cataracts, ARMD, and conductive and sensory hearing loss can be diagnosed early. Many people now have their eyes and hearing tested by high-street optometrists in addition to relying on GP referrals.

People who have diabetes and hypertension need tests more often because both conditions can adversely affect sight and hearing. People with a family history of glaucoma should also be encouraged to undergo regular testing because this condition (which is not part of the normal ageing process) can be hereditary.

Bishop PN et al (2004) Age-related changes on the surface of vitreous collagen fibrils. Investigative Ophthalmology and Visual Science; 45: 4, 1041-1046.

Bitsios P et al (1996) Changes in autonomic function with age: a study of pupillary kinetics in healthy young and old people. Age and Ageing; 25: 6, 432-438.

Boyd K (2016) What is Presbyopia? American Academy of Ophthalmology.

Danner CJ, Harris JP (2003) Hearing loss and the aging ear. Geriatrics and Aging; 6: 5, 40-43.

Dry Eye WorkShop (2007) The definition and classification of dry eye disease: report of the Definition and Classification Subcommittee of the International Dry Eye WorkShop. Ocular Surface; 5: 2, 75-92.

Forrester JV et al (2001) The Eye: Basic Sciences in Practice. New York, NY: Elsevier.

Garrity J (2016) Entropion and Ectropion.

Horak FB (2006) Postural orientation and equilibrium: what do we need to know about neural control of balance to prevent falls? Age and Ageing; 35: Suppl 2, ii7ii11.

Howarth A, Shone GR (2006) Ageing and the auditory system. Postgraduate Medical Journal; 82: 965, 166-171.

Knight J et al (2017) Diabetes management 2: long-term complications due to poor control. Nursing Times; 113: 4, 45-48.

Kujawa SG, Liberman MC (2006) Acceleration of age-related hearing loss by early noise exposure: evidence of a misspent youth. Journal of Neuroscience; 26: 7, 2115-2123.

Liang L et al (2011) Ocular surface morbidity in eyes with senile sunken upper eyelids. Ophthalmology; 118: 12, 2487-2492.

Liu TC, Chen YS (2000) Aging and external ear resonance. International Journal of Audiology; 39: 5, 235-237.

Parham K et al (2011) Challenges and opportunities in presbycusis. Otolaryngology Head and Neck Surgery; 144: 4, 491-495.

Rauch SD et al (2001) Decreasing hair cell counts in aging humans. Annals of the New York Academy of Sciences; 942: 220-227.

Rukmini AV et al (2017) Pupillary responses to short-wavelength light are preserved in aging. Scientific Reports; 7: 43832.

Shupert C, Horak F (2017) Balance and Aging. Portland, OR: Vestibular Disorders Association.

Sobel RK, Tienor B (2013) The coming age of enophthalmos. Current Opinion in Ophthalmology; 24: 5, 500-505.

Sogebi OA (2015) Middle ear impedance studies in elderly patients: implications on age-related hearing loss. Brazilian Journal of Otorhinolaryngology; 81: 2, 133-140.

Truscott RJW (2003) Human cataract: the mechanisms responsible; light and butterfly eyes. International Journal of Biochemistry and Cell Biology; 35: 11, 1500-1504.

Turner PL, Mainster MA (2008) Circadian photoreception: ageing and the eyes important role in systemic health. British Journal of Ophthalmology; 92: 11, 1439-1444.

Weinstein BE (2000) Geriatric Audiology. New York, NY: Thieme.

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Anatomy and physiology of ageing 6: the eyes and ears - Nursing Times

CARBONDALE Howard H. Olson passed away peacefully with his wife and children by his side, in Murrells Inlet, South Carolina, on June 21, 2017.

A frequent and enthusiastic traveler, he had just completed his last of many trips. This one was, to celebrate his 90th birthday and the graduation of his youngest grandchild in California.

Howard Olson was born May 23, 1927, in Chicago, the son of the late Halfdan G. Olson and Ruth Swanson Olson.

Howard was a lifelong learner. He graduated from Carl Schurz High School in Chicago in 1944. Upon graduating, he enrolled at the University of Wisconsin, receiving a B.S. in dairy science in 1948. He continued his education at the University of Minnesota, earning his doctorate in dairy science and physiology in 1952. His interest in agriculture stemmed from spending his summers on his mothers parents farm in Amery, Wisconsin.

He met his first wife, Maurine Fay Schroeder, while they were studying at the University of Minnesota. They were married in 1951.

After receiving his doctorate, Howard served as director of research with Curtiss Candy Farms in Cary, for two years before leaving to become a professor of Dairy Science at Southern Illinois University in Carbondale, where he remained until his retirement in 1989. In addition to teaching multiple generations of students, he taught and conducted research programs in Dairy Cattle Management and Reproductive Physiology.

Howards interest in dairy science as well as his willingness to embrace new experiences and challenges led to a lifetime of teaching, research, and learning throughout the world. He served as director of International Agriculture at SIU, managing programs in Brazil, Zambia, and Pakistan. His interest in international work started in 1966 when he, family in tow, left for Cairo, Egypt, where he took a position as a Fulbright Lecturer at Ain Shams University for a year. Once this interest started, it never abated. Shortly after his time in Egypt, he took off for Hyderabad, India, serving as a Dairy Cattle Management trainer for Peace Corps volunteers. Then, in 1981, he served as a Fulbright lecturer at the University of Peradeniya in Kandy, Sri Lanka. His international interest outlasted his tenure as a professor. After retirement, he served on Farmer-to-Farmer assignments in Egypt, Kazakhstan, Armenia, Nicaragua, and Mexico.

His international forays aside, Howard lived in Carbondale, for most of his professional life, where he and Maurine raised their four children. After Maurines death, he moved to Chapel Hill, North Carolina, where his daughter Kirstens family lived at the time in 1992. In 2001, he married Ethel Devendorf and moved to Murrells Inlet, South Carolina. During the last two decades of his life, he traveled the world with Ethel and enjoyed summers every year on Stoner Lake in Caroga Lake, New York.

He was a member of Alpha Gamma Rho Social Fraternity and of Alpha Zeta and Sigma Xi Honorary Fraternities.

Throughout his life, he was an active member of the Lutheran Church.

He was predeceased by his first wife, Maurine Fay Schroeder Olson (1990) and his daughter, Gwen Lucas (1999).

He is survived by his wife, Ethel Irene Klinger Devendorf; his children, Kurt Olson and family of Miami, Florida, Kirsten Olson and family of Arlington, Virginia, and Karin Olson and family of Palo Alto, California; and his seven grandchildren; two great-grandsons; three stepchildren; eight stepgrandchildren; and 13 great-stepgrandchildren.

Memorials may be presented to Shepherd of the Sea Lutheran Church, (www.shepherdofthesea.com).

An interment service will be conducted at the Sunset Memorial Cemetery in Minneapolis, Minnesota.

Burroughs Funeral Home and Cremation Services (843-651-1440) of Murrells Inlet is assisting the family.

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Howard H. Olson - The Southern