Chapter 14: Homeostasis

Homeostasis
- Homeostasis: Maintenance of a relatively constant internal environment for the cells within
the body
• Temperature – controls rate of metabolic reactions & enzyme activity
• Water potential – controls osmotic movement
• Concentration of glucose – controls gradients
• pH – controls enzyme efficiency
• Blood pressure
• O2, CO2, NH3, etc.

- Homeostatic control
+) Components of a homeostatic system

• Stimulus: change in external / internal environment which triggers a response

• Receptor: cell / tissue sensitive to a specific stimulus & communicates with a control
center by generating nerve impulses (nervous) or sending chemical messengers (endocrine)
• Effector: A tissue / organ that carries out an action in response to a stimulus, e.g. muscles,
glands
• Corrective action: the response / series of responses that return a physiological factor to
the set point
+) Most control mechanisms use a negative feedback control loop: a process in which a change in a
parameter brings about counteracting processes which return it to normal

• The parameter fluctuates around a set point: the ideal value of a physiological factor that
the body controls in homeostasis
• The control mechanism can only the parameter in a certain range around the set point

*Positive feedback: A change in a parameter brings about processes that move its level further in
the direction of the initial change

• The parameter grows exponentially, then stops completely at a certain limit

• Examples: accumulation of platelets during blood clotting, production of oxytocin during
childbirth, etc.
 The kidney & kidney structure

*Assimilation: The conversion of molecules from food in to other molecules that the body needs

- Excretion
+) Excretion: the removal of toxic / metabolic waste products from the body
+) Deamination: the breakdown of excess amino acids in the liver, by the removal of the amine
group

• The amine group (-NH2) and an extra H form NH3

• The keto acid may be respired, or stored as glycogen / fat

+) Ammonia is soluble & toxic

• Increases blood pH → messes up metabolic processes & cell signaling
• NH3 is converted to urea (less soluble & toxic) through the urea cycle
2NH3 + CO2 → CO(NH2)2 + H2O
urea

+) Urea is the main nitrogenous excretory product of humans

• There are also small amounts of others e.g. creatinine, uric acid
o Creatine is produced in the liver from amino acids & used in the muscles as an
energy store, but can also be converted to creatinine to be excreted
o Uric acid is produced from breakdown of purines from nucleotides

- General kidney structure

+) 3 main parts
• Cortex - under the kidneys
• Medulla - central area
• Renal pelvis – where the ureter joins

+) IG recall – general components & keywords

• Afferent arteriole: arteriole leading to glomerular capillaries
• Efferent arteriole: arteriole leading away from glomerular capillaries, into the nephron’s
capillary network
• The blood flows into venules & collect at (a branch of) the renal vein at the end
- Production of urine is a 2-step process:
• Ultrafiltration: filtration on a molecular scale, separating small molecules from larger ones,
such as proteins
o Small molecules, including urea, are filtered out of the blood in the glomerulus, into
the Bowman’s capsule, forming filtrate
• Selective reabsorption: movement of certain (useful) substances from the filtrate in
nephrons back into the blood
 Ultrafiltration & selective reabsorption
*Nephron structures & functions
Structure Functions Adaptations
Glomerulus & Bowman’s - Ultrafiltration of blood by size - Podocytes wrap around
capsule - Basement membrane acts as capillaries
a filter - Basement membrane
- Afferent arteriole wider than
efferent arteriole to increase
capillary pressure
- Mass of capillaries
Proximal convoluted tubule - Selective reabsorption - Thin layer of cells
- Active transport of glucose & - Microvilli providing large SA
amino acids - Many mitochondria in cells
- Osmosis of some water
- Diffusion of some salts
Loop of Henle - Reabsorption of water - Thick and thin sections
- Descending limb is water - Thick limbs have cuboidal
permeable epithelial cells
- Ascending limb is
impermeable but actively
pumps out salt
- Creates a salt gradient in the
medulla

Distal convoluted tubule

Collecting duct

- Ultrafiltration
+) There are 3 layers between the capillary blood & the Bowman’s capsule’s lumen:
• Cell layer - Capillary endothelium, perforated by many circular holes, 60-80 nm in diameter
(bigger than normal capillaries)
• Basement membrane, made up of a network of collagen & glycoproteins; also acts as a
filter
• Cell layer – Podocytes, epithelial cells making up the inner lining of the BM’s lumen; have
many tiny finger-like projections with gaps in between

+) The basement membrane acts as the filter

• Stops proteins with a relative molecular mass of over 69000 from passing through &
escaping the glomerular capillaries
o Includes RBCs, WBCs, platelets
➔ Composition of glomerular filtrate is nearly identical to blood plasma, just without the plasma
proteins

*Rate of glomerular filtration

+) Rate of filtration in human kidneys: ~125 cm3min-1
+) The heart supplies the kidney with over a litre of blood every minute

+) The high rate of diffusion is due to the water potential gradient

+) 2 components to the water potential gradient:
• Pressure
o Diameter of blood vessels decrease from the afferent to efferent arterioles ➔
glomerular capillaries have higher pressure ➔ increases water potential inside
capillaries ➔ blood to filtrate potential
• Solute concentration
o Plasma proteins cannot be filtered out ➔ lower water potential

• Overall, the effect of pressure outweighs the effect of concentration ➔ water moves down
the gradient from the blood into the capsule

- Selective reabsorption
+) Proximal convoluted tubule
• Where most of the reabsorption takes place
• The PCT lining is made of a single layer of cuboidal epithelial cells, adapted to the
reabsorption function:
o Many microvilli on the surface facing the lumen to increase SA
o Many co-transporter proteins in the luminal membrane
o Tight junctions that hold adjacent cells tightly, to prevent passage of fluids between
cells
o Many mitochondria to provide energy for Na+ - K+ pumps in basal (basement)
membranes
+) Reabsorption of sodium
• Na+ is moved from inside the PCT (cuboidal epithelial) cell into the blood by Na+ - K+ pumps

+) Co-transport & reabsorption of glucose

• Lower Na+ concentration ➔ Na+ passively diffuses from the filtrate (inside the lumen)
through the luminal membrane, into the PCT cell
• This diffusion is only allowed through Na+ co-transporter proteins in the luminal membrane,
which co-transports glucose / certain amino acids, after which is actively transported
through the basal membrane into blood
o Glucose & amino acids are completely reabsorbed
o Vitamins, Na+, Cl- is also reabsorbed (not completely)
o This is considered secondary active transport, as energy was used in pumping Na +
ions into the blood, which facilitated the passive movement & co-transport

+) Reabsorption of water
• The removal of solutes from the filtrate increases its water potential ➔ Water moves down
gradient into the blood

+) Other shit
• The CSM is partially permeable to urea ➔ Urea is also reabsorbed – diffuses down
concentration gradient into blood
• Reabsorption greatly reduces filtrate volume
- Reabsorption in the loop of Henle & collecting duct
+) The loops create a very high Na+ / Cl- ion concentration in the medulla’s tissue fluid, lowering the
water potential
• This is partly achieved by active Na+ - Cl- symporters in the thick ascending limb (see pic)
• There is both passive & active movement

+) Water is reabsorbed by osmosis down ψ gradient

• Functionally, to prevent dehydration

+) This reabsorption continues at the collecting duct (running into medulla)

• Water moves until ψurine = ψmedulla
o Urine concentration between organisms vary according to degree of salts pumped
into tissue fluid
• The degree of this absorption is controlled by the antidiuretic hormone (ADH)
o ADH: hormone secreted by the posterior pituitary gland; increases water
reabsorption in the kidneys (and consequently reduces water loss in urine)

- Reabsorption in the distal convoluted tubule:

• The first half functions like the ascending limb of the loop of Henle
• The second half functions like the collecting duct
o In both, Na+ - K+ pumps actively transport Na+ / K+ into tissue fluid / tubule

Control of water content – collecting duct

- Osmoregulation: the control of water potential of blood and tissue fluid by controlling the
water content and/or ion concentration (particularly Na )+

• Involves the hypothalamus, posterior pituitary gland & kidneys

o Hypothalamus: a gland located in the brain that monitors several factors: blood
water potential, and also body temperature, heart rate, etc.

- Blood water potential is monitored by osmoreceptors in the hypothalamus

• Sends nerve impulses along neurons that terminate at the PPG when blood water potential
falls below a set point
• ADH is then produced by the PPG, which enter the blood & increases reabsorption of water
in the kidneys
 (overview)

*Osmoreceptors
• When osmotic pressure of blood rises, water moves down the osmotic gradient into the cell
➔ swelling
o Reverse occurs in low osmotic pressure
• Triggers neuronal signals to be sent to the hypothalamus

- ADH action
• ADH targets the luminal membrane of the collecting duct cells, increasing their
permeability to water

- Mechanism
 • The change in permeability is due to an increase in number of aquaporins – water-
permeable membranes - in the luminal membrane

• ADH binds to receptor proteins, which stimulates production of cyclic AMP (cAMP), a
second messenger
o cAMP activates a signaling cascade, leading to phosphorylation of aquaporin
o This activation causes the aquaporin-containing vesicles to move towards and fuse
with the membrane → increased permeability
• The cells contain ready-made vesicles with membranes containing aquaporins, which
become part of the luminal membrane when the vesicle fuses with it
• Water now moves through aquaporins, out of the tubule & into tissue fluid (note that the
medulla still has a much lower water potential)

- Reverse process
• When there is excess water, osmoreceptors in hypothalamus aren’t stimulated ➔ PPG
stops secreting ADH
• No ADH stimulation ➔ aquaporins move back into the cell as vesicles
• This reverse process is not instant as it takes time for ADH to break down

Intracellular signaling
- Steroid hormones
• Lipophilic – can freely diffuse across cell membranes
• Binds to receptors in the cytoplasm / nucleus of the target cell → forms a complex that
moves into the nucleus, binds directly to DNA & activates gene expression
 • Slow-acting, but carries out permanent changes
• E.g. Estrogen, progesterone, testosterone

- Peptide hormones
• Hydrophilic & lipophobic – can’t cross cell membranes freely
• Binds to surface receptors (usually GPCRs), activating a signal transduction pathway
(carried by internal intermediaries)
o This enables amplification of the signal

• Rapid acting
• E.g. Insulin, glucagon, leptin, ADH, oxytocin

- Mechanism
+) First messenger: G-protein
• GPCRs (G-protein-coupled receptors): a family of CSM receptors; transmits signals inside
the cell through a G-protein; has 7 different protein segments that crosses the CSM
• G-proteins (Guanine nucleotide-binding proteins): a family of proteins; acts as molecular
switches inside cells; involved in transmitting signals from a variety of stimuli outside a cell
to its interior
o This signal transfer is usually in the form of transfer of phosphate groups

+) Intracellular signaling
• Phosphorylation is the main method of signaling inside cells
• Transfer of phosphate group is catalyzed by kinase
o Cells contain different kinases with different targets
• Common signaling pathways: MAPK, Ca2+ ions, cAMP, inositol phosphates

+) Second messenger: cAMP (cyclic AMP)

 o

• The G-protein phosphorylates & activates adenylyl cyclase to convert ATP → cAMP
o Removes 2 phosphates & links the leftover phosphate to the sugar → cyclic
o cAMP: a small molecule made from ATP; acts as a second messenger in the
cytoplasm
• cAMP activates PKA (protein kinase A) to perform phosphorylation & continue the signal
o cAMP is found in many cell types, with different target proteins in each → allows
cAMP to produce different responses in different contexts
 The control of blood glucose

- Homeostatic control of glucose is controlled by 2 hormones, secreted by endocrine tissue in the

pancreas
• Islets of Langerhans: groups of cells that make up the endocrine tissue; secretes glucagon
& insulin
• The islets are made up of:
o α cells, secretes glucagon
▪ Glucagon: small peptide hormone, brings about decrease of glucose
concentration
o β cells, secretes insulin
▪ Insulin: small peptide hormone, brings about increase of glucose
concentration
• α and β cells act as receptors & the central control for the mechanism
• Insulin & glucagon acts as coordinators

- Control mechanism

*Insulin secretion
 • At high glucose concentration, glucose enters B cells by facilitated diffusion
• The cells respire the glucose → ATP
• High concentrations of ATP cause potassium channels in the B cell’s CSM to close,
producing a change in the membrane potential
• The change in membrane potential causes the voltage-gated calcium channels to open
• In response to the Ca2+ influx, insulin-containing vesicles move towards the CSM and go out
by exocytosis

- Response to increase in blood glucose

• α and β cells detect the increase in glucose conc.
• α cells stop secreting glucagon, and β cells start secreting insulin, which is carried around
the whole body by blood
 • Insulin binds to a CSM receptor (being a protein, can’t pass CSM)
• The binding of insulin activates movement of vesicles containing GLUT4 towards the CSM to
fuse with it & increase the cell’s permeability to glucose
o Activation
▪ GCPR activates G-protein → G-protein activation → adenylyl cyclase
activation → cAMP production → protein kinase A activation → … → vesicle
activated

o Glucose only enters cells by facilitated diffusion through GLUT transporter proteins
o Several types of GLUT
▪ GLUT4, found in muscle cells (shown above)
▪ GLUT1, found in brain cells & GLUT2, found in liver cells; these are always on
the CSM and aren’t affected by insulin

• Insulin also stimulates activation of glucokinase, which phosphorylates glucose to trap it

inside
o Also begins glycolysis & respiration

*Glycogenesis
• Insulin also stimulates activation of phosphofructokinase & glycogen synthase
• These 2 enzymes together catalyze glycogenesis – synthesis of glycogen by addition of
glucose monomers
- Response to increase in blood glucose
• After detection, α cells start secreting glucagon, and β cells stop secreting insulin

• Glucagon binds to receptors in the CSM of liver cells

o Muscle cells don’t have glucagon receptors
• The binding causes a conformational change in the receptor protein, activating a G-protein
• The G-protein activates adenylyl cyclase – an enzyme that catalyzes the conversion of ATP
to cAMP, a second messenger (explained above)

• An enzyme cascade commences, amplifying the signal:

o cAMP binds to & activates protein kinase A
o PKA adds phosphate groups to & activates phosphorylase kinase
o Phosphorylase kinase adds phosphate groups to & activates glycogen
phosphorylase

• Finally, glycogen phosphorylase catalyzes glycogenolysis: breakdown of glycogen →

glucose by removal of glucose monomers
• The glucose diffuses out through GLUT2 proteins

*Extra info
➢ Glucagon also stimulates gluconeogenesis: formation of glucose from non-carb sources,
e.g. amino acids, fatty acids, glycerol, pyruvate, lactate, etc.
➢ Blood glucose conc. is never perfectly constant, due to time delays in cell signaling &
corrective actions
➢ Adrenaline can also increase blood glucose conc.
 o Can activate the same enzyme cascade as glucagon (identical action)
o Stimulates breakdown of glycogen in muscle cells for local use

*Glycolysis – glycogenesis – glycogenolysis – gluconeogenesis

• Glycolysis: breakdown of glucose into pyruvate; the first step in aerobic respiration
• Gluconeogenesis: formation of glucose from pyruvate (& also other shit)
• Glycogenesis: formation of glycogen by addition of glucose (monomers)
• Glycogenolysis: breakdown of glycogen into glucose monomers

- Diabetes

- Measuring blood glucose concentration

+) High blood glucose conc. means inability to reabsorb all the glucose, and may indicate diabetes

+) Urine testing
• Test strips can be used to test urine for many factors e.g. glucose, pH, protein, etc.
• Contains immobilized glucose oxidase & peroxidase, which triggers 2 reactions together

o Oxidized chromogen is brown

o Higher glucose concentration → darker color
 • Covered by a cellulose membrane to only allow glucose passage
• Only shows concentration at the time of urine collection, not in present time

+) Biosensor
• A device that uses a biological material e.g. enzymes, to measure concentration of a
chemical compound
• Uses the same immobilized enzyme – recognition layer structure (with a few extra steps),
but can directly measure from blood & give a reading quickly
 Homeostasis in plants

- Guard cells: kidney-shaped epidermal cells, found in pairs surrounding a stoma & controlling its
opening – closure

- Typical features of guard cells:

• Thick cell walls that face the outside air & the stomatal pore
o Outer wall has thick waxy cuticle
o Inner walls are thinner
• Cellulose microfibrils arranged into bands around cell
• Absence of plasmodesmata in cell walls
• Folded CSM & contains many proteins
• High chloroplast & mitochondria density in cytoplasm
o Mitochondria have many cristae
• Chloroplasts have thylakoids, but fewer grana (than mesophyll cells); starch grains enlarge /
shrink during the night / day
• Nucleus occupies a larger proportion of the cell
• Several small vacuoles instead of 1 big one

- Opening and closing of stomata

• Stomata have daily rhythms of opening & closing
• Stomata can respond to changes in environmental conditions by opening & closing
o Regulation of stomatal aperture balances the need for CO2 uptake by diffusion with
the need to minimize water loss by transpiration

• Opening response:
o Increased light intensity
o Low CO2 concentrations in air spaces

• Closing response:
o Darkness (low LI)
o High CO2 concentrations in air spaces
o Low humidity
o High temperature
o Water stress – limited water supply from roots / high rate of transpiration

- Opening & closing mechanism

• Guard cells open when gaining water & becoming turgid, and close when losing water &
becoming flaccid

+) Opening
• In response to light, ATP-powered proton pumps in the guard cell CSM transport H+ out
• Decrease in H+ causes CSM protein channels to open and K+ to move inside
o K+ movement is due to both a charge gradient & concentration gradient
 ➔ Electrochemical gradient: gradient across CSM that involves both an ion conc.
gradient & potential difference
o Other ions also enter to help maintain charge balance e.g. Cl-, NO3-
• Increased solute concentration from entry of K+ decreases the guard cell’s water potential
• Water moves in through aquaporins into the vacuole, expanding it
• Turgor pressure increases ➔ stoma opens
o During this, starch is broken down into negative malate ions that enter the vacuole,
which maintain electrical balance

• Guard cells have unevenly thickened walls

o Gets thicker getting closer to the pore (pore-adjacent walls are thickest)
• Cellulose microfibrils
o Forces the guard cells to lengthen instead of expanding
o Due to the outer walls being thinner, the guard cells push on each other when turgid
+) Closing
• H+ ion closes & K+ leaves to enter neighboring cells
• Malate ions return to the chloroplast & turn back into starch
• Water potential increases & water leaves

*Effects of closing stoma

• Reduces CO2 uptake & rate of transpiration
o Transpiration is important, for cooling & maintaining the transpiration stream that
supplies water & minerals
• Stoma closure only occurs in conditions of water stress
• Abscisic acid (ABA): an inhibitory plant growth regulator that causes closure of stomata
during dry conditions

- Mechanism of ABA / abscisic acid

• Can bind with ABA receptors on guard cells to inhibit proton pumps
• Can also stimulate Ca2+ movement into the cell cytoplasm through the CSM & tonoplast
• Role of Ca2+
o Activates protein channels to allow negative ions to leave the cell, which in turn
stimulates opening of more protein channels, which allows K+ to leave
o Stimulates closure of protein channels that allow K+ to enter