William-Adolphe Bouguereau (1825-1905) - Thirst (1886)

Thirst is the craving for potable fluids, resulting in the basic instinct of animals to drink. It is an essential mechanism involved in

fluid balance Fluid balance is an aspect of the homeostasis of organisms in which the amount of water in the organism needs to be controlled, via osmoregulation and behavior, such that the concentrations of electrolytes ( salts in solution) in the various b ...

. It arises from a lack of fluids or an increase in the concentration of certain

osmolite Osmolytes are low-molecular weight organic compounds that influence the properties of biological fluids. Their primary role is to maintain the integrity of cells by affecting the viscosity, melting point, and ionic strength of the aqueous solution. ...

s, such as

sodium Sodium is a chemical element with the symbol Na (from Latin ''natrium'') and atomic number 11. It is a soft, silvery-white, highly reactive metal. Sodium is an alkali metal, being in group 1 of the periodic table. Its only stable ...

. If the water volume of the body falls below a certain threshold or the osmolite concentration becomes too high, structures in the

brain A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a ve ...

detect changes in blood constituents and signal thirst. Continuous

dehydration In physiology, dehydration is a lack of total body water, with an accompanying disruption of metabolic processes. It occurs when free water loss exceeds free water intake, usually due to exercise, disease, or high environmental temperature. Mil ...

can cause acute and chronic diseases, but is most often associated with

renal The kidneys are two reddish-brown bean-shaped organs found in vertebrates. They are located on the left and right in the retroperitoneal space, and in adult humans are about in length. They receive blood from the paired renal arteries; bloo ...

and neurological disorders. Excessive thirst, called

polydipsia Polydipsia is excessive thirst or excess drinking.Porth, C. M. (1990). ''Pathophysiology: Concepts of altered health states''. Philadelphia: J.B. Lippincott Company. The word derives from the Greek () "very thirsty", which is derived from (, "mu ...

, along with excessive urination, known as

polyuria Polyuria () is excessive or an abnormally large production or passage of urine (greater than 2.5 L or 3 L over 24 hours in adults). Increased production and passage of urine may also be termed diuresis. Polyuria often appears in conjunction wit ...

, may be an indication of

diabetes mellitus Diabetes, also known as diabetes mellitus, is a group of metabolic disorders characterized by a high blood sugar level ( hyperglycemia) over a prolonged period of time. Symptoms often include frequent urination, increased thirst and increased ...

diabetes insipidus Diabetes insipidus (DI), recently renamed to Arginine Vasopressin Deficiency (AVP-D) and Arginine Vasopressin Resistance (AVP-R), is a condition characterized by large amounts of dilute urine and increased thirst. The amount of urine produced ...

. There are

receptors Receptor may refer to: *Sensory receptor, in physiology, any structure which, on receiving environmental stimuli, produces an informative nerve impulse *Receptor (biochemistry), in biochemistry, a protein molecule that receives and responds to a n ...

and other systems in the body that detect a decreased volume or an increased osmolite concentration. Some sources distinguish "extracellular thirst" from "intracellular thirst", where extracellular thirst is thirst generated by decreased volume and intracellular thirst is thirst generated by increased osmolite concentration.

Detection

It is vital for organisms to be able to maintain their fluid levels in very narrow ranges. The goal is to keep the interstitial fluid, the fluid outside the cell, at the same concentration as the intracellular fluid, the fluid inside the cell. This condition is called isotonic and occurs when the same levels of solutes are present on either side of the

cell membrane The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment ( ...

so that the net water movement is zero. If the interstitial fluid has a higher concentration of solutes (or a lower concentration of water) than the intracellular fluid, it will pull water out of the cell. This condition is called

hypertonic In chemical biology, tonicity is a measure of the effective osmotic pressure gradient; the water potential of two solutions separated by a partially-permeable cell membrane. Tonicity depends on the relative concentration of selective membrane- ...

and if enough water leaves the cell, it will not be able to perform essential chemical functions. The animal will then become thirsty in response to the demand for water in the cell. After the animal drinks water, the interstitial fluid becomes less concentrated of solutes (more concentrated of water) than the intracellular fluid and the cell will fill with water as it tries to equalize the concentrations. This condition is called hypotonic and can be dangerous because it can cause the cell to swell and rupture. One set of receptors responsible for thirst detects the concentration of interstitial fluid. The other set of receptors detects blood volume.

Decreased volume

This is one of two types of thirst and is defined as thirst caused by loss of blood volume (hypovolemia) without depleting the intracellular fluid. This can be caused by blood loss, vomiting, and diarrhea. This loss of volume is problematic because if the total blood volume falls too low the heart cannot circulate blood effectively and the eventual result is hypovolemic shock. The vascular system responds by constricting blood vessels thereby creating a smaller volume for the blood to fill. This mechanical solution, however, has definite limits and usually must be supplemented with increased volume. The loss of blood volume is detected by cells in the kidneys and triggers thirst for both water and salt via the renin-angiotensin system.

Renin-angiotensin system

Hypovolemia leads to activation of the

renin angiotensin system Renin (etymology and pronunciation), also known as an angiotensinogenase, is an aspartic protease protein and enzyme secreted by the kidneys that participates in the body's renin–angiotensin–aldosterone system (RAAS)—also known as the r ...

(RAS) and is detected by cells in the kidney. When these cells detect decreased blood flow due to the low volume they secrete an enzyme called

renin Renin (etymology and pronunciation), also known as an angiotensinogenase, is an aspartic protease protein and enzyme secreted by the kidneys that participates in the body's renin–angiotensin–aldosterone system (RAAS)—also known as the r ...

. Renin then enters the blood where it catalyzes a protein called

angiotensinogen Angiotensin is a peptide hormone that causes vasoconstriction and an increase in blood pressure. It is part of the renin–angiotensin system, which regulates blood pressure. Angiotensin also stimulates the release of aldosterone from the a ...

angiotensin I Angiotensin is a peptide hormone that causes vasoconstriction and an increase in blood pressure. It is part of the renin–angiotensin system, which regulates blood pressure. Angiotensin also stimulates the release of aldosterone from the a ...

. Angiotensin I is then almost immediately converted by an enzyme already present in the blood to the active form of the protein, angiotensin II. Angiotensin II then travels in the blood until it reaches the

posterior pituitary gland The posterior pituitary (or neurohypophysis) is the posterior lobe of the pituitary gland which is part of the endocrine system. The posterior pituitary is not glandular as is the anterior pituitary. Instead, it is largely a collection of axona ...

and the adrenal cortex, where it causes a cascade effect of hormones that cause the kidneys to retain water and sodium, increasing blood pressure. It is also responsible for the initiation of

drinking Drinking is the act of ingesting water or other liquids into the body through the mouth, proboscis, or elsewhere. Humans drink by swallowing, completed by peristalsis in the esophagus. The physiological processes of drinking vary widely am ...

behavior and salt appetite via the

subfornical organ The subfornical organ (SFO) is one of the circumventricular organs of the brain. Its name comes from its location on the ventral surface of the fornix near the interventricular foramina (foramina of Monro), which interconnect the lateral ventr ...

Others

* Arterial baroreceptors sense a decreased arterial pressure, and signal to the

central nervous system The central nervous system (CNS) is the part of the nervous system consisting primarily of the brain and spinal cord. The CNS is so named because the brain integrates the received information and coordinates and influences the activity of all p ...

in the

area postrema The area postrema, a paired structure in the medulla oblongata of the brainstem, is a circumventricular organ having permeable capillaries and sensory neurons that enable its dual role to detect circulating chemical messengers in the blood and t ...

and

nucleus tractus solitarii In the human brainstem, the solitary nucleus, also called nucleus of the solitary tract, nucleus solitarius, and nucleus tractus solitarii, (SN or NTS) is a series of purely sensory nuclei (clusters of nerve cell bodies) forming a vertical column ...

. *

Cardiopulmonary receptor Baroreceptors (or archaically, pressoreceptors) are sensors located in the carotid sinus (at the bifurcation of External carotid artery, external and Internal carotid artery, internal carotids) and in the aortic arch. They sense the blood pressure ...

s sense a decreased blood volume, and signal to

and

Cellular dehydration and osmoreceptor stimulation

Osmometric thirst occurs when the solute concentration of the interstitial fluid increases. This increase draws water out of the cells, and they shrink in volume. The solute concentration of the interstitial fluid increases by high intake of sodium in diet or by the drop in volume of extracellular fluids (such as

blood plasma Blood plasma is a light amber-colored liquid component of blood in which blood cells are absent, but contains proteins and other constituents of whole blood in suspension. It makes up about 55% of the body's total blood volume. It is the intr ...

and

cerebrospinal fluid Cerebrospinal fluid (CSF) is a clear, colorless body fluid found within the tissue that surrounds the brain and spinal cord of all vertebrates. CSF is produced by specialised ependymal cells in the choroid plexus of the ventricles of the ...

) due to loss of water through perspiration, respiration, urination and defecation. The increase in interstitial fluid solute concentration causes water to migrate from the cells of the body, through their membranes, to the extracellular compartment, by

osmosis Osmosis (, ) is the spontaneous net movement or diffusion of solvent molecules through a selectively-permeable membrane from a region of high water potential (region of lower solute concentration) to a region of low water potential (region o ...

, thus causing cellular dehydration. Clusters of cells (

osmoreceptors An osmoreceptor is a sensory receptor primarily found in the hypothalamus of most homeothermic organisms that detects changes in osmotic pressure. Osmoreceptors can be found in several structures, including two of the circumventricular organs – t ...

) in the organum vasculosum of the lamina terminalis (OVLT) and

(SFO), which lie outside of the blood brain barrier can detect the concentration of blood plasma and the presence of angiotensin II in the blood. They can then activate the

median preoptic nucleus The median preoptic nucleus is located dorsal to the other three nuclei of the preoptic area of the anterior hypothalamus. The hypothalamus is located just beneath the thalamus, the main sensory relay station of the nervous system, and is con ...

which initiates water seeking and ingestive behavior. Destruction of this part of the hypothalamus in humans and other animals results in partial or total loss of desire to drink even with extremely high salt concentration in the extracellular fluids. Page 872 In addition, there are

viscera In biology, an organ is a collection of tissues joined in a structural unit to serve a common function. In the hierarchy of life, an organ lies between tissue and an organ system. Tissues are formed from same type cells to act together in a f ...

osmoreceptor An osmoreceptor is a sensory receptor primarily found in the hypothalamus of most homeothermic organisms that detects changes in osmotic pressure. Osmoreceptors can be found in several structures, including two of the circumventricular organs – ...

s which project to the

and

in the brain.

Salt craving

Because sodium is also lost from the plasma in hypovolemia, the body's need for salt proportionately increases in addition to thirst in such cases. This is also a result of the renin-angiotensin system activation.

Elderly

In adults over the age of 50 years, the body's thirst sensation reduces and continues diminishing with age, putting this population at increased risk of

. Several studies have demonstrated that elderly persons have lower total water intakes than younger adults, and that women are particularly at risk of too low an intake. In 2009, the

European Food Safety Authority The European Food Safety Authority (EFSA) is the agency of the European Union (EU) that provides independent scientific advice and communicates on existing and emerging risks associated with the food chain. EFSA was established in February 2002, ...

(EFSA) included water as a macronutrient in its dietary reference values for the first time."EFSA Panel on Dietetic Products, Nutrition, and Allergies (NDA)". ''EFSA Journal'' 2010;8(3):1459. Recommended intake volumes in the elderly are the same as for younger adults (2.0 L/day for females and 2.5 L/day for males) as despite lower energy consumption, the water requirement of this group is increased due to a reduction in renal concentrating capacity.IoM (Institute of Medicine), 2004. Dietary Reference Intakes for Water, Potassium, Sodium, Chloride, and Sulfate. National Academies Press, Washington, D.C.

Thirst quenching

According to preliminary research, quenching of thirst – the

homeostatic In biology, homeostasis (British also homoeostasis) (/hɒmɪə(ʊ)ˈsteɪsɪs/) is the state of steady internal, physical, and chemical conditions maintained by living systems. This is the condition of optimal functioning for the organism and ...

mechanism to stop drinking – occurs via two neural phases: a "preabsorptive" phase which signals quenched thirst many minutes before fluid is absorbed from the stomach and distributed to the body via the circulation, and a "postabsorptive" phase which is regulated by brain structures sensing to terminate fluid ingestion. The preabsorptive phase relies on sensory inputs in the mouth,

pharynx The pharynx (plural: pharynges) is the part of the throat behind the mouth and nasal cavity, and above the oesophagus and trachea (the tubes going down to the stomach and the lungs). It is found in vertebrates and invertebrates, though its st ...

esophagus The esophagus ( American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to ...

, and upper gastrointestinal tract to anticipate the amount of fluid needed, providing rapid signals to the brain to terminate drinking when the assessed amount has been consumed. The postabsorptive phase occurs via blood monitoring for

osmolality Molality is a measure of the number of moles of solute in a solution corresponding to 1 kg or 1000 g of solvent. This contrasts with the definition of molarity which is based on a specified volume of solution. A commonly used unit for molali ...

, fluid volume, and sodium balance, which are collectively sensed in brain

circumventricular organs Circumventricular organs (CVOs) ( circum-: around ; ventricular: of ventricle) are structures in the brain characterized by their extensive and highly permeable capillaries, unlike those in the rest of the brain where there exists a blood� ...

linked via neural networks to terminate thirst when fluid balance is established. Thirst quenching varies among animal species, with dogs, camels, sheep, goats, and deer replacing fluid deficits quickly when water is available, whereas humans and horses may need hours to restore fluid balance.

Neurophysiology

The areas of the brain that contribute to the sense of thirst are mainly located in the

midbrain The midbrain or mesencephalon is the forward-most portion of the brainstem and is associated with vision, hearing, motor control, sleep and wakefulness, arousal (alertness), and temperature regulation. The name comes from the Greek ''mesos'', " ...

and the

hindbrain The hindbrain or rhombencephalon or lower brain is a developmental categorization of portions of the central nervous system in vertebrates. It includes the medulla, pons, and cerebellum. Together they support vital bodily processes. Metencephal ...

. Specifically, the

hypothalamus The hypothalamus () is a part of the brain that contains a number of small nuclei with a variety of functions. One of the most important functions is to link the nervous system to the endocrine system via the pituitary gland. The hypothalamu ...

appears to play a key role in the regulation of thirst. The

and

signal to the subfornical organ and to the

lateral parabrachial nucleus The parabrachial nuclei, also known as the parabrachial complex, are a group of nucleus (neuroanatomy), nuclei in the dorsolateral pons that surrounds the superior cerebellar peduncle as it enters the brainstem from the cerebellum. They are named ...

. The latter signaling relies on the neurotransmitter serotonin. The signal from the lateral parabrachial nucleus is relayed to the

. The median preoptic nucleus and the subfornical organ receive signals of decreased volume and increased osmolite concentration. Finally, the signals are received in

cortex Cortex or cortical may refer to: Biology * Cortex (anatomy), the outermost layer of an organ ** Cerebral cortex, the outer layer of the vertebrate cerebrum, part of which is the ''forebrain'' *** Motor cortex, the regions of the cerebral cortex i ...

areas of the

forebrain In the anatomy of the brain of vertebrates, the forebrain or prosencephalon is the rostral (forward-most) portion of the brain. The forebrain (prosencephalon), the midbrain (mesencephalon), and hindbrain (rhombencephalon) are the three primary ...

where thirst arises. The

and the organum vasculosum of the lamina terminalis contribute to regulating the overall bodily

by signalling to the hypothalamus to form

vasopressin Human vasopressin, also called antidiuretic hormone (ADH), arginine vasopressin (AVP) or argipressin, is a hormone synthesized from the AVP gene as a peptide prohormone in neurons in the hypothalamus, and is converted to AVP. It then trave ...

, which is later released by the

pituitary gland In vertebrate anatomy, the pituitary gland, or hypophysis, is an endocrine gland, about the size of a chickpea and weighing, on average, in humans. It is a protrusion off the bottom of the hypothalamus at the base of the brain. The ...