CAMPBELL BIOLOGY IN FOCUS © 2014 Pearson Education, Inc. Urry Cain Wasserman Minorsky Jackson Reece Lecture Presentations by Kathleen Fitzpatrick and Nicole Tunbridge 32 Homeostasis and Endocrine Signaling

© 2014 Pearson Education, Inc. Overview: Diverse Forms, Common Challenges  Anatomy is the study of the biological form of an organism  Physiology is the study of the biological functions an organism performs  Form and function are closely correlated

© 2014 Pearson Education, Inc. Figure 32.1

© 2014 Pearson Education, Inc. Concept 32.1: Feedback control maintains the internal environment in many animals  Multicellularity allows for cellular specialization with particular cells devoted to specific activities  Specialization requires organization and results in an internal environment that differs from the external environment

© 2014 Pearson Education, Inc. Hierarchical Organization of Animal Bodies  Cells form a functional animal body through emergent properties that arise from levels of structural and functional organization  Cells are organized into  Tissues, groups of cells with similar appearance and common function  Organs, different types of tissues organized into functional units  Organ systems, groups of organs that work together

© 2014 Pearson Education, Inc. Table 32.1

© 2014 Pearson Education, Inc.  The specialized, complex organ systems of animals are built from a limited set of cell and tissue types  Animal tissues can be grouped into four categories  Epithelial  Connective  Muscle  Nervous

© 2014 Pearson Education, Inc. Figure 32.2 Axons of neurons Skeletal muscle tissue Blood vessel Loose connective tissue Blood Epithelial tissue Collagenous fiber Epithelial tissue Lumen 10  m Basal surface Apical surface Nervous tissue Glia 20  m Plasma White blood cells (Confocal LM) 50  m Red blood cells 100  m Elastic fiber Nuclei Muscle cell 100  m

© 2014 Pearson Education, Inc. Regulating and Conforming  Faced with environmental fluctuations, animals manage their internal environment by either regulating or conforming  An animal that is a regulator uses internal mechanisms to control internal change despite external fluctuation  An animal that is a conformer allows its internal condition to change in accordance with external changes

© 2014 Pearson Education, Inc. Figure 32.3 River otter (temperature regulator) 40 Largemouth bass (temperature conformer) Ambient (environmental) temperature (  C) Body temperature (  C)

© 2014 Pearson Education, Inc.  An animal may regulate some internal conditions and not others  For example, a fish may conform to surrounding temperature in the water, but it regulates solute concentrations in its blood and interstitial fluid (the fluid surrounding body cells)

© 2014 Pearson Education, Inc. Homeostasis  Organisms use homeostasis to maintain a “steady state” or internal balance regardless of external environment  In humans, body temperature, blood pH, and glucose concentration are each maintained at a constant level  Regulation of room temperature by a thermostat is analogous to homeostasis

© 2014 Pearson Education, Inc. Figure 32.4 Sensor/ control center: Thermostat turns heater off. Sensor/ control center: Thermostat turns heater on. Stimulus: Room temperature increases. Stimulus: Room temperature decreases. Room temperature increases. Room temperature decreases. Set point: Room temperature at 20  C Response: Heating stops. Response: Heating starts.

© 2014 Pearson Education, Inc.  Animals achieve homeostasis by maintaining a variable at or near a particular value, or set point  Fluctuations above or below the set point serve as a stimulus; these are detected by a sensor and trigger a response  The response returns the variable to the set point

© 2014 Pearson Education, Inc.  Homeostasis in animals relies largely on negative feedback, an output that that reduces the stimulus  Homeostasis moderates, but does not eliminate, changes in the internal environment  Set points and normal ranges for homeostasis are usually stable, but certain regulated changes in the internal environment are essential

© 2014 Pearson Education, Inc. Thermoregulation: A Closer Look  Thermoregulation is the process by which animals maintain an internal temperature within a tolerable range  Endothermic animals generate heat by metabolism; birds and mammals are endotherms  Ectothermic animals gain heat from external sources; ectotherms include most invertebrates, fishes, amphibians, and nonavian reptiles  regulate temperature by behavioral means

© 2014 Pearson Education, Inc. Figure 32.8a Sensor/control center: Thermostat in hypothalamus Body temperature decreases. Homeostasis: Internal body temperature of approximately 36–38  C Response: Blood vessels in skin dilate. Stimulus: Increased body temperature Response: Sweat

© 2014 Pearson Education, Inc. Figure 32.8b Stimulus: Decreased body temperature Body temperature increases. Homeostasis: Internal body temperature of approximately 36–38  C Response: Shivering Sensor/control center: Thermostat in hypothalamus Response: Blood vessels in skin constrict.

© 2014 Pearson Education, Inc. Figure 32.8 Sensor/control center: Thermostat in hypothalamus Stimulus: Decreased body temperature Body temperature increases. Body temperature decreases. Homeostasis: Internal body temperature of approximately 36–38  C Response: Blood vessels in skin dilate. Response: Shivering Sensor/control center: Thermostat in hypothalamus Response: Blood vessels in skin constrict. Stimulus: Increased body temperature Response: Sweat

© 2014 Pearson Education, Inc. Figure 32.5 (a) A walrus, an endotherm (b) A lizard, an ectotherm

© 2014 Pearson Education, Inc. Figure 32.6 Radiation Convection Evaporation Conduction

© 2014 Pearson Education, Inc.  In response to changes in environmental temperature, animals can alter blood (and heat) flow between their body core and their skin  Vasodilation, the widening of the diameter of superficial blood vessels, promotes heat loss  Vasoconstriction, the narrowing of the diameter of superficial blood vessels, reduces heat loss Circulatory Adaptations for Thermoregulation

© 2014 Pearson Education, Inc.  The arrangement of blood vessels in many marine mammals and birds allows for countercurrent exchange  Countercurrent heat exchangers transfer heat between fluids flowing in opposite directions and reduce heat loss

© 2014 Pearson Education, Inc. Figure 32.7 Canada goose Blood flow Artery Vein Heat transfer Cool blood Warm blood Key 30  35  C 99 20  10  18  27  33  231

© 2014 Pearson Education, Inc. Physiological Thermostats and Fever  Thermoregulation in mammals is controlled by a region of the brain called the hypothalamus  The hypothalamus triggers heat loss or heat- generating mechanisms  Fever is the result of a change to the set point for a biological thermostat

© 2014 Pearson Education, Inc. Concept 32.2: Endocrine signals trigger homeostatic mechanisms in target tissues There are two major systems for controlling and coordinating responses to stimuli: the endocrine and nervous systems  In the endocrine system, signaling molecules released into the bloodstream by endocrine cells reach all locations in the body  In the nervous system, neurons transmit signals along dedicated routes, connecting specific locations in the body

© 2014 Pearson Education, Inc.  Signaling molecules sent out by the endocrine system are called hormones  Hormones may have effects in a single location or throughout the body  Only cells with receptors for a certain hormone can respond to it  The endocrine system is well adapted for coordinating gradual changes that affect the entire body

© 2014 Pearson Education, Inc. Figure 32.9 Cell body of neuron Response Hormone Nerve impulse Signal travels everywhere. Signal travels to a specific location. Response Stimulus Nerve impulse Blood vessel Endocrine cell (a) Signaling by hormones Axons Axon (b) Signaling by neurons

© 2014 Pearson Education, Inc. Figure 32.9a Cell body of neuron Hormone Signal travels everywhere. Signal travels to a specific location. Stimulus Nerve impulse Blood vessel Endocrine cell (a) Signaling by hormones Axon (b) Signaling by neurons

© 2014 Pearson Education, Inc.  In the nervous system, signals called nerve impulses travel along communication lines consisting mainly of axons  Other neurons, muscle cells, and endocrine and exocrine cells can all receive nerve impulses  Nervous system communication usually involves more than one type of signal  The nervous system is well suited for directing immediate and rapid responses to the environment

© 2014 Pearson Education, Inc. Figure 32.9b Response Nerve impulse Response Blood vessel (a) Signaling by hormones Axons (b) Signaling by neurons

© 2014 Pearson Education, Inc. Simple Endocrine Pathways  Digestive juices in the stomach are extremely acidic and must be neutralized before the remaining steps of digestion take place  Coordination of pH control in the duodenum relies on an endocrine pathway

© 2014 Pearson Education, Inc. Figure Response Hormone Low pH in duodenum S cells of duodenum secrete the hormone secretin ( ). Pancreas Stimulus Blood vessel Endocrine cell Pathway Example Bicarbonate release Target cells Negative feedback 

© 2014 Pearson Education, Inc.  The release of acidic stomach contents into the duodenum stimulates endocrine cells there to secrete the hormone secretin  This causes target cells in the pancreas to raise the pH in the duodenum  The pancreas can act as an exocrine gland, secreting substances through a duct, or as an endocrine gland, secreting hormones directly into interstitial fluid

© 2014 Pearson Education, Inc. Neuroendocrine Pathways  Hormone pathways that respond to stimuli from the external environment rely on a sensor in the nervous system  In vertebrates, the hypothalamus integrates endocrine and nervous systems  Signals from the hypothalamus travel to a gland located at its base, called the pituitary gland

© 2014 Pearson Education, Inc. Figure 32.11a Pancreas Insulin Glucagon Testes (in males) Androgens Parathyroid glands Parathyroid hormone (PTH) Ovaries (in females) Estrogens Progestins Thyroid gland Thyroid hormone (T 3 and T 4 ) Calcitonin Pineal gland Melatonin Major Endocrine Glands and Their Hormones Hypothalamus Pituitary gland Anterior pituitary Posterior pituitary Oxytocin Vasopressin (antidiuretic hormone, ADH) Adrenal glands (atop kidneys) Adrenal medulla Epinephrine and norepinephrine Adrenal cortex Glucocorticoids Mineralocorticoids

© 2014 Pearson Education, Inc. Figure 32.11b Posterior pituitary Neurosecretory cells of the hypothalamus Hypothalamic hormones Hypothalamus Testes or ovaries Portal vessels Pituitary hormones Anterior pituitary Endocrine cells Thyroid gland Adrenal cortex Mammary glands MelanocytesLiver, bones, other tissues HORMONE TARGET Prolactin FSHTSH ACTH MSH GH

© 2014 Pearson Education, Inc. Figure 32.11c TSH circulation throughout body Anterior pituitary Thyroid gland Stimulus Response Thyroid hormone Thyroid hormone circulation throughout body TSH − TRH Neurosecretory cell Negative feedback Hypothalamus − Sensory neuron

© 2014 Pearson Education, Inc. Figure 32.11d Thyroid scan Low level of iodine uptake High level of iodine uptake

© 2014 Pearson Education, Inc. Feedback Regulation in Endocrine Pathways  A feedback loop links the response back to the original stimulus in an endocrine pathway  While negative feedback dampens a stimulus, positive feedback reinforces a stimulus to increase the response

© 2014 Pearson Education, Inc. Pathways of Water-Soluble and Lipid-Soluble Hormones  The hormones discussed thus far are proteins that bind to cell-surface receptors and that trigger events leading to a cellular response  The intracellular response is called signal transduction  A signal transduction pathway typically has multiple steps

© 2014 Pearson Education, Inc.  Lipid-soluble hormones have receptors inside cells  When bound by the hormone, the hormone- receptor complex moves into the nucleus  There, the receptor alters transcription of particular genes

© 2014 Pearson Education, Inc. Multiple Effects of Hormones  Many hormones elicit more than one type of response  For example, epinephrine is secreted by the adrenal glands and can raise blood glucose levels, increase blood flow to muscles, and decrease blood flow to the digestive system  Target cells vary in their response to a hormone because they differ in their receptor types or in the molecules that produce the response

© 2014 Pearson Education, Inc. Figure Different cellular responses Same receptors but different intracellular proteins (not shown) Glycogen breaks down and glucose is released from cell. Different cellular responses Different receptors Glycogen deposits Vessel dilates. Vessel constricts. (c) Intestinal blood vessel (b) Skeletal muscle blood vessel (a) Liver cell Epinephrine  receptor Epinephrine  receptor Epinephrine  receptor

© 2014 Pearson Education, Inc. Evolution of Hormone Function  Over the course of evolution the function of a given hormone may diverge between species  For example, thyroid hormone plays a role in metabolism across many lineages, but in frogs it has taken on a unique function: stimulating the resorption of the tadpole tail during metamorphosis  Prolactin also has a broad range of activities in vertebrates

© 2014 Pearson Education, Inc. Figure Tadpole Adult frog

© 2014 Pearson Education, Inc. Concept 32.3: A shared system mediates osmoregulation and excretion in many animals  Osmoregulation is the general term for the processes by which animals control solute concentrations in the interstitial fluid and balance water gain and loss

© 2014 Pearson Education, Inc. Osmosis and Osmolarity  Cells require a balance between uptake and loss of water  Osmolarity, the solute concentration of a solution, determines the movement of water across a selectively permeable membrane  If two solutions are isoosmotic, the movement of water is equal in both directions  If two solutions differ in osmolarity, the net flow of water is from the hypoosmotic to the hyperosmotic solution

© 2014 Pearson Education, Inc. Osmoregulatory Challenges and Mechanisms  Osmoconformers, consisting of some marine animals, are isoosmotic with their surroundings and do not regulate their osmolarity  Osmoregulators expend energy to control water uptake and loss in a hyperosmotic or hypoosmotic environment

© 2014 Pearson Education, Inc.  Marine and freshwater organisms have opposite challenges  Marine fish drink large amounts of seawater to balance water loss and excrete salt through their gills and kidneys  Freshwater fish drink almost no water and replenish salts through eating; some also replenish salts by uptake across the gills

© 2014 Pearson Education, Inc. Figure 32.15a Gain of water and salt ions from food SALT WATER Gain of water and salt ions from drinking seawater Excretion of salt ions from gills Excretion of salt ions and small amounts of water in scanty urine from kidneys Osmotic water loss through gills and other parts of body surface Salt Water (a) Osmoregulation in a marine fish

© 2014 Pearson Education, Inc. Figure 32.15b Gain of water and some ions in food FRESH WATER Uptake of salt ions by gills Excretion of salt ions and large amounts of water in dilute urine from kidneys Osmotic water gain through gills and other parts of body surface Salt Water (b) Osmoregulation in a freshwater fish

© 2014 Pearson Education, Inc. Nitrogenous Wastes  The type and quantity of an animal’s waste products may greatly affect its water balance  Among the most significant wastes are nitrogenous breakdown products of proteins and nucleic acids  Some animals convert toxic ammonia (NH 3 ) to less toxic compounds prior to excretion

© 2014 Pearson Education, Inc. Figure Most aquatic animals, including most bony fishes ProteinsNucleic acids Amino acids Nitrogenous bases Amino groups Mammals, most amphibians, sharks, some bony fishes Many reptiles (including birds), insects, land snails Ammonia Urea Uric acid

© 2014 Pearson Education, Inc. Figure 32.16a Most aquatic animals, including most bony fishes Mammals, most amphibians, sharks, some bony fishes Many reptiles (including birds), insects, land snails Ammonia Urea Uric acid

© 2014 Pearson Education, Inc.  Ammonia excretion is most common in aquatic organisms  Vertebrates excrete urea, a conversion product of ammonia, which is much less toxic  Insects, land snails, and many reptiles including birds excrete uric acid as a semisolid paste  It is less toxic than ammonia and generates very little water loss, but it is energetically more expensive to produce than urea

© 2014 Pearson Education, Inc. Excretory Processes  In most animals, osmoregulation and metabolic waste disposal rely on transport epithelia  These layers of epithelial cells are specialized for moving solutes in controlled amounts in specific directions

© 2014 Pearson Education, Inc.  Many animal species produce urine by refining a filtrate derived from body fluids  Key functions of most excretory systems  Filtration: Filtering of body fluids  Reabsorption: Reclaiming valuable solutes  Secretion: Adding nonessential solutes and wastes from the body fluids to the filtrate  Excretion: Releasing processed filtrate containing nitrogenous wastes from the body

© 2014 Pearson Education, Inc. Figure Capillary Filtration Excretory tubule Filtrate Reabsorption Secretion Urine Excretion

© 2014 Pearson Education, Inc. Figure 32.19bc Nephron Organization Collecting duct Branch of renal vein Vasa recta Efferent arteriole from glomerulus Distal tubule Afferent arteriole from renal artery Glomerulus Bowman’s capsule Proximal tubule Peritubular capillaries Descending limb Ascending limb Loop of Henle

© 2014 Pearson Education, Inc. Concept 32.4: Hormonal circuits link kidney function, water balance, and blood pressure  The capillaries and specialized cells of Bowman’s capsule are permeable to water and small solutes but not blood cells or large molecules  The filtrate produced there contains salts, glucose, amino acids, vitamins, nitrogenous wastes, and other small molecules

© 2014 Pearson Education, Inc. Figure Filtrate OUTER MEDULLA H2OH2O Salts (NaCI and others) HCO 3 − Glucose, amino acids HH Some drugs Passive transport Active transport Key INNER MEDULLA CORTEX Descending limb of loop of Henle H2OH2O Interstitial fluid NH 3 HH Nutrients HCO 3 − KK NaCI Proximal tubule H2OH2O Thick segment of ascending limb HH Urea HCO 3 − KK NaCI Distal tubule H2OH2O Thin segment of ascending limb NaCI H2OH2O Collecting duct Urea

© 2014 Pearson Education, Inc. From Blood Filtrate to Urine: A Closer Look Proximal tubule  Reabsorption of ions, water, and nutrients takes place in the proximal tubule  Molecules are transported actively and passively from the filtrate into the interstitial fluid and then capillaries  Some toxic materials are actively secreted into the filtrate

© 2014 Pearson Education, Inc. Descending limb of the loop of Henle  Reabsorption of water continues through channels formed by aquaporin proteins  Movement is driven by the high osmolarity of the interstitial fluid, which is hyperosmotic to the filtrate  The filtrate becomes increasingly concentrated all along its journey down the descending limb

© 2014 Pearson Education, Inc. Ascending limb of the loop of Henle  The ascending limb has a transport epithelium that lacks water channels  Here, salt but not water is able to move from the tubule into the interstitial fluid  The filtrate becomes increasingly dilute as it moves up to the cortex

© 2014 Pearson Education, Inc. Distal tubule  The distal tubule regulates the K + and NaCl concentrations of body fluids  The controlled movement of ions contributes to pH regulation

© 2014 Pearson Education, Inc. Collecting duct  The collecting duct carries filtrate through the medulla to the renal pelvis  Most of the water and nearly all sugars, amino acids, vitamins, and other nutrients are reabsorbed into the blood  Urine is hyperosmotic to body fluids

© 2014 Pearson Education, Inc. Figure Passive transport Active transport mOsm/L CORTEX OUTER MEDULLA INNER MEDULLA 300 H2OH2O 1, H2OH2O H2OH2O H2OH2O H2OH2O H2OH2O H2OH2O ,200

© 2014 Pearson Education, Inc. Figure Passive transport Active transport mOsm/L CORTEX OUTER MEDULLA INNER MEDULLA 300 H2OH2O 1, H2OH2O H2OH2O H2OH2O H2OH2O H2OH2O H2OH2O NaCI ,200

© 2014 Pearson Education, Inc. Figure Passive transport Active transport H2OH2O mOsm/L CORTEX Urea OUTER MEDULLA INNER MEDULLA 300 NaCI H2OH2O H2OH2O 1, H2OH2O H2OH2O H2OH2O H2OH2O H2OH2O H2OH2O NaCI , ,200 Urea NaCI H2OH2O H2OH2O H2OH2O H2OH2O H2OH2O

© 2014 Pearson Education, Inc.  The loop of Henle and surrounding capillaries act as a type of countercurrent system  This system involves active transport and thus an expenditure of energy  Such a system is called a countercurrent multiplier system

© 2014 Pearson Education, Inc.  The filtrate in the ascending limb of the loop of Henle is hypoosmotic to the body fluids by the time it reaches the distal tubule  The filtrate descends to the collecting duct, which is permeable to water but not to salt  Osmosis extracts water from the filtrate to concentrate salts, urea, and other solutes in the filtrate