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Chapter 17: Excretory System

Recommended Article : 【Biology】 Table of Contents for Biology


1. Regulation of Ingestion

2. Nitrogenous Wastes

3. Types of Excretory Systems

4. Structure of Kidneys

5. Kidney Functions

6. Kidney Regulation

7. Kidney Disorders

8. Sweat



1. Regulation of Ingestion



2. Nitrogenous Wastes

⑴ Generation of Nitrogenous Wastes

① Amino Acid Breakdown : Reaction in which amino groups are removed during amino acid metabolism, producing ammonia

○ Liver combines ammonia with CO2 via the ornithine cycle to synthesize urea

○ Mitochondria and cytoplasm are involved

○ Urea formation occurs in the cytoplasm

○ Kidneys secrete ammonia into renal tubules

○ Amino Group Transfer Reaction (Glutamine, Asparagine) : Reaction transferring amino groups to form 2-oxoacids


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Figure 1. Ornithine Cycle


② Nucleic Acid Breakdown : Production of uric acid

○ Humans lack enzymes for uric acid breakdown

○ 25% excretion via digestive system, 75% excretion via kidneys

○ Accumulation of uric acid can lead to gout and uric acid kidney stones (composed of uric acid, phosphates, calcium, oxalates, etc.)

○ Gout : Main causes include genetics, kidney disorders, medications, diabetes, alcohol consumption, etc.

⑵ Characteristics of Nitrogenous Wastes and Animal Physiology


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Figure 2. Types of Nitrogenous Wastes


① Ammonia (NH3, ammonia)

○ Characteristics : High solubility, high toxicity (because ammonia readily accepts H+ ions)

○ (During excretion) Water requirement : 1 mL per 1 g of nitrogen

○ Relevant animals : Mostly cartilaginous fish, aquatic invertebrates

② Urea

○ Characteristics : Much weaker toxicity compared to ammonia (10,000 times), excretable in concentrated form

○ Energy required for urea synthesis

○ (During excretion) Water requirement : 1 mL per 50 g of nitrogen

○ Relevant animals : Mostly mammals, amphibians, cartilaginous fish, some bony fish

③ Uric Acid

○ Characteristics : Non-toxic, low solubility (semi-solid state), minimal water consumption during excretion (water-saving effect)

○ Requires more energy than urea synthesis

○ (During excretion) Water requirement : 1 mL per 500 g of nitrogen

○ Relevant animals : Birds, reptiles, terrestrial arthropods

⑶ Evolution and Environmental Impact on Nitrogenous Wastes

① Methods of waste elimination while preventing water loss : Urea, uric acid

② Determined by Reproductive Strategy

○ Eggs of shell-less amphibians, embryos of mammals : Preferentially excrete soluble waste (ammonia becomes more toxic upon accumulation)

○ Eggs of birds, reptiles with shells : Excrete uric acid (non-toxic even upon accumulation)

③ Influenced by Habitat

○ Example : Terrestrial turtles (uric acid), aquatic turtles (urea, ammonia)

④ Endotherms have more nitrogenous waste compared to ectotherms



3. Types of Excretory Systems

⑴ Protonephridia (Flame Cell System) : Excretory system of flatworms

① Protonephridia branch like tree branches, with flame cells (composed of cap cells and flame cells) at the terminal end

○ Flame cells have a flame-like appearance, giving them their name

② Cell-to-cell filtration occurs through a membrane connecting interstitial fluid and flame cells

⑵ Metanephridia : Excretory system of annelids responsible for osmoregulation and waste excretion

① 1st Filtrate enters metanephridium through tubules surrounded by cilia

② 2nd Tubule cells alter the composition of the filtrate as it flows through the tubules

③ 3rd Diluted urine is expelled outside through the excretory pore

⑶ Malpighian Tubules : Excretory system of terrestrial arthropods

① 1st Salts, water, and nitrogenous wastes enter Malpighian tubules

② 2nd Malpighian tubules excrete urine into the hindgut

③ 3rd Feces and urine are expelled through the anus

⑷ Kidneys : Excretory system of vertebrates



4. Structure of Kidneys

⑴ Pathway of Urine Formation : Nephron → Collecting Duct Renal Papilla Ureter Bladder Urethra

① Renal Cortex : Nephron

② Renal Medulla : Henle’s loop, collecting duct, renal papilla

③ Related Blood Vessels : Afferent arteriole → Peritubular capillaries → Efferent arteriole

⑵ Nephron

① Functional unit of kidneys, each kidney has 1.25 million nephrons (totaling 145 km), composed of renal corpuscle and renal tubule

② Renal Corpuscle (Malpighian Body) : Collectively refers to glomerulus and Bowman’s capsule, located in the renal cortex

○ Glomerulus : A cluster of capillaries

○ Bowman’s Capsule : Double-layered capsule surrounding the glomerulus

③ Renal Tubule

○ Macula Densa : Cells surrounding the renal tubule, prevents free movement of ions by tight junctions

○ Proximal Convoluted Tubule (PCT) : Present in the cortex. Close to the afferent arteriole

○ Henle’s Loop : Present in the medulla. Found only in mammals and birds

○ Distal Convoluted Tubule (DCT) : Present in the cortex. Close to the efferent arteriole

④ Cortical Nephron, Juxtamedullary Nephron

○ Nephrons span both the renal cortex and medulla

○ Cortical Nephron : Nephron with a bias toward the cortex

○ Juxtamedullary Nephron : Nephron with a bias toward the medulla

⑤ Intercalated Cells : Present in the DCT and collecting duct

○ Type A Intercalated Cells : Active during acidosis. Secrete H+ into the lumen. Reabsorb HCO3-. Reabsorb K+

○ Type B Intercalated Cells : Active during alkalosis. Reabsorb H+. Secrete HCO3-. Secrete K+



5. Kidney Functions

⑴ 20-25% of cardiac output (500-600 mL/minute) passes through the afferent arterioles

⑵ Filtration, Reabsorption, Secretion, Excretion

① Filtration : Small molecules (H2O, salts, etc.) are non-selectively filtered in the glomerulus

② Reabsorption : Tubule cells transport useful substances from the filtrate back to the body fluids

③ Secretion : Toxic substances and excess ions from body fluids (capillaries) are secreted into the filtrate (tubules)

④ Excretion : Filtrate leaves the kidneys and is expelled from the body

⑶ 1st. Filtration

① Glomerular Filtration Pressure : Driving force for filtration

○ Glomerular Filtration Pressure = Glomerular Capillary Pressure (Blood Pressure) - (Bowman’s Capsule Osmotic Pressure + Plasma Colloid Osmotic Pressure) = 55 - (30 + 15) = 10 (mmHg)

○ Plasma Colloid Osmotic Pressure : Caused by proteins in the blood that pull water back into the blood

○ Bowman’s Capsule Osmotic Pressure : Counteracts the hydrostatic pressure generated in Bowman’s capsule

② Difference in blood flow between afferent and efferent arterioles determines filtration rate

○ Afferent arterioles are larger in diameter compared to efferent arterioles

○ Constriction of afferent arterioles : Decreases blood flow to glomerular capillaries, reducing glomerular capillary pressure Decreased filtration rate

○ Constriction of efferent arterioles : Limits blood outflow, increasing glomerular capillary pressure Increased filtration rate

○ Dilation of afferent arterioles : Increases blood flow to glomerular capillaries, raising glomerular capillary pressure Increased filtration rate

○ Dilation of efferent arterioles : Facilitates blood outflow, decreasing glomerular capillary pressure Decreased filtration rate

○ Vasoconstriction : Increases pressure in the arterial portion, decreases in the venous portion

③ Filtration Rate

○ Renal Blood Flow (RBF) (unit: mL/min) : Blood flow through renal vessels in 1 minute

○ Renal Plasma Flow (RPF) (unit: mL/min) : Plasma flow through renal vessels in 1 minute

○ Renal Threshold : Concentration at which a substance starts to be secreted

○ Hematocrit = Blood Hematocrit = Plasma / Blood = RPF / RBF

PAH : Completely removed from plasma

Creatine : Not reabsorbed or secreted in the renal tubules

○ Produced by breakdown of creatine phosphate in muscles at a fairly constant rate

○ Cleared by kidneys, frequently used to measure GFR

○ Used as a safety marker for kidney function

Negative Pressure: No secretion and reabsorption through the nephron

○ Chart for Filtration Rate Calculation: Calculation of filtration rate can be understood through the principle of mass conservation.


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Table 1. Filtration Rate Calculation Chart


○ Higher filtration rate of positive ions due to the negative charge of the Bowman’s capsule


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Figure 3. Influence of Size and Charge of Substances on Filtration


④ Regulation of GFR (Glomerular Filtration Rate)

○ Afferent Arteriole: Constriction decreases RPF (Renal Plasma Flow) and GFR, Dilation increases RPF and GFR

○ Efferent Arteriole: Constriction decreases RPF, increases GFR, Dilation increases RPF, decreases GFR

⑤ Glomerular Filtration Barrier


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Figure 4. Glomerular Filtration Barrier


Component 1: Capillary endothelial cell: Refers to fenestrae or pores, allowing the passage of substances smaller than 70 nm.

Component 2: Glomerular basement membrane: High collagen density, negatively charged, allows passage of substances smaller than 6-6.5 nm.

Component 3: Podocyte extension: Thin membrane connecting podocytes, with a width of approximately 4-11 nm.

⑥ Changes in GFR with Age


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Table 2. Changes in GFR with Age


⑦ Classification of CKD (Chronic Kidney Disease)


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Table 3. Classification of CKD

(Source: CKD Work Group, 2013)


⑷ 2nd. Concentration of Filtrate

① Model of Concentration of Filtrate

○ Two-solute Model: Maintains high osmotic pressure of inner renal medulla due to NaCl and solutes.

○ Countercurrent Multiplier: Consumes energy to establish concentration gradient.

② Glomerulus → Proximal Convoluted Tubule → Descending Limb of Henle → Ascending Limb of Henle → Distal Convoluted Tubule → Collecting Duct

○ Reabsorption of 75% of filtered fluid (67% NaCl, H2O)

○ Na+ Reabsorption (Active transport, Na+/K+ pump) → Cl- reabsorption (Passive transport) for charge balance

○ H2O Reabsorption (Passive transport)

○ Aquaporin-mediated water movement is facilitated diffusion, one type of passive transport, in response to Na+ reabsorption.

○ HCO3- Reabsorption: Na+-linked secondary active transport, contributes to acid-base balance in body fluids.

○ K+ Reabsorption: Passive transport, followed by conditional secretion in the distal convoluted tubule.

○ Secondary Active Transport of Hydrophilic Nutrients

○ Related to the threshold for secretion

○ Threshold (Plasma threshold): Concentration at which a substance starts being secreted

○ Not related to contraction of afferent and efferent arterioles

○ 100% Reabsorption of Glucose: Na+-linked secondary active transport. 320 mg glucose/minute, using glucose transporter GluT2.

○ 100% Reabsorption of Amino Acids: Na+-linked secondary active transport.

○ Secretion: NH3, solutes, creatinine, drugs, etc.

○ pH Regulation: H+ secretion


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Figure 5. Renal Threshold


③ Glomerulus → Proximal Convoluted Tubule → Descending Limb of Henle → Ascending Limb of Henle → Distal Convoluted Tubule → Collecting Duct

○ Descending Limb of Henle: Impermeable to water, permeable to NaCl. AQP-1 is expressed.

○ Water exchange between Descending Limb of Henle and renal medulla

○ Osmolarity comparison: Filtrate < Renal medulla

○ Filtrate gets concentrated as water is reabsorbed from Descending Limb of Henle to renal medulla.

○ Without countercurrent multiplication, renal medulla remains diluted.

○ Water exchange between Straight Vasa Recta and renal medulla

○ Return of osmolarity: Osmolarity increases in descending Straight Vasa Recta, then decreases as it ascends from the bottom of Henle’s loop, leading to water reabsorption.

○ High Na+ concentration in descending Straight Vasa Recta leads to high water reabsorption, increasing blood pressure.

④ Glomerulus → Proximal Convoluted Tubule → Descending Limb of Henle → Ascending Limb of Henle → Distal Convoluted Tubule → Collecting Duct

○ Impermeable to water, permeable to NaCl NaCl reabsorption occurs

○ Tight junctions prevent water reabsorption.

○ Thin Ascending Limb: Reabsorption of Na+ only, passive transport

○ Thick Ascending Limb: Reabsorption of Na+ and Cl- together, active transport [Countercurrent multiplication 1]

⑤ Glomerulus → Proximal Convoluted Tubule → Descending Limb of Henle → Ascending Limb of Henle → Distal Convoluted Tubule → Collecting Duct

○ Variable substance transport due to hormonal regulation, mostly active transport

○ Na+ pump regulated by RAAS (Renin-Angiotensin-Aldosterone System)

○ 1st. Decreased water → Decreased GFR

○ 2nd. Slower filtrate generation → Increased Na+ reabsorption in thick Ascending Limb

○ 3rd. Dilute urine stimulates JGA (Juxtaglomerular Apparatus)

○ 4th. JGA stimulates granular cells

○ 5th. Granular cells secrete renin

○ 6th. Renin activates angiotensinogen to angiotensin I in the liver

○ 7th. Angiotensin I is converted to angiotensin II by angiotensin-converting enzyme (ACE)

○ 8th. Mechanisms to conserve water, including aldosterone, are activated

○ Na+-K+ Pump: Moves 3 Na+ ions out and 2 K+ ions into cells

○ Na+-K+-Cl- co-transporter in Ascending Limb follows Na+ concentration gradient, leading to Na+, K+, and Cl- reabsorption in the lumen of the tubule.

○ Na+ reabsorption (Active transport) → Cl- reabsorption (Passive transport) for charge balance

○ Conditional expression of aquaporin channels for water transport

○ H2O reabsorption (Passive transport)

○ Active transport of HCO3- for pH regulation: Reabsorption of bicarbonate ions

○ K+ secretion: Active transport to regulate filtrate osmolarity

○ Selective secretion: NH3, solutes, creatinine, drugs, etc.

○ pH Regulation: H+ secretion for pH control

⑥ Glomerulus → Proximal Convoluted Tubule → Descending Limb of Henle → Ascending Limb of Henle → Distal Convoluted Tubule → Collecting Duct

○ Na+ pump regulated by RAAS

○ 1st. Decreased water → Decreased GFR

○ 2nd. Slower filtrate generation → Increased Na+ reabsorption in thick Ascending Limb

○ 3rd. Dilute urine stimulates JGA

○ 4th. JGA stimulates granular cells

○ 5th. Granular cells secrete renin

○ 6th. Renin activates angiotensinogen to angiotensin I in the liver

○ 7th. Angiotensin I is converted to angiotensin II by angiotensin-converting enzyme (ACE)

○ 8th. Mechanisms to conserve water, including aldosterone, are activated

○ Na+-K+ Pump: Moves 3 Na+ ions out and 2 K+ ions into cells

○ Na+-K+-Cl- co-transporter in Ascending Limb follows Na+ concentration gradient, leading to Na+, K+, and Cl- reabsorption in the lumen of the tubule.

○ Na+ reabsorption (Active transport) → Cl- reabsorption (Passive transport) for charge balance

○ Conditional expression of aquaporin channels for water transport

○ H2O reabsorption (Passive transport)

○ Central diabetes insipidus: Problem in hypothalamus → Normal with ADH administration

○ Nephrogenic diabetes insipidus: Problem with ADH receptors

○ Acid-base balance

○ Type A Intercalated Cells: ↑ H+ → ↑ K+ → Hyperkalemia

○ Type B Intercalated Cells: ↓ H+ → ↓ K+ → Hypokalemia

○ Changes in H+ and K+ move together

○ Reabsorption of solutes: Continuous storage of solutes in renal medulla to increase medullary osmolarity [Countercurrent multiplication 2]

○ Some of the stored solutes in renal medulla move into the filtrate of Henle’s loop

⑦ Changes in Osmolarity of Adjacent Vasa Recta (Peritubular Capillaries)

○ Blood in descending vessels has higher osmolarity

○ Blood in ascending vessels has lower osmolarity

○ Therefore, the change in osmolarity of blood passing through the kidney is nearly 0

⑸ 3rd. Excretion

① Urine excretion: 1.5 liters/day

○ Bladder can store up to 500 ml, releases 200-300 ml of urine at a time

○ Reabsorption from filtrate reduces volume

② Excretion of 10 g of salt per day (= 99.5% reabsorption of salt)


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Figure 6. Function of Nephron



6. Regulation of the Kidney

⑴ Antidiuretic Hormone (ADH): Also known as vasopressin

① Produced in hypothalamus, stored in posterior pituitary, and released

② Function 1: Osmoregulation

Step 1: Increase in osmolarity

Step 2: ADH secretion triggered by osmoreceptors in hypothalamus

Step 3: ADH secretion

Step 4: Increased expression of aquaporins in Distal Convoluted Tubule and Collecting Duct

Step 5: Increased water reabsorption

○ Step 6: Osmoregulation (300 mOsm/L)

○ Osmoreceptors stimulate thirst, inducing drinking behavior

③ Function 2: Vasoconstriction of arterioles due to smooth muscle contraction → Increased blood pressure

④ Mechanism of increased water reabsorption by ADH

○ 1st. ADH binds to the collecting duct.

○ 2nd. Receptor stimulates the cAMP second messenger pathway.

○ 3rd. Insertion of aquaporins, including aquaporin, into the membrane on the side of the urinary bladder.

○ 4th. Efficient reabsorption of water from the urinary bladder due to aquaporins.

⑤ Causes of polyuria

○ Causes : Defective ADH receptors or inadequate production/secretion of ADH ×

○ Insufficient aquaporins leading to difficulty in water reabsorption.

⑥ Caffeine, alcohol promote diuresis by inhibiting the secretion of antidiuretic hormone (ADH).

⑦ ADH contributes more to plasma sodium concentration homeostasis than the RAAS (Renin-Angiotensin-Aldosterone System).

⑵ Renin-Angiotensin-Aldosterone System (RAAS)

① Juxtaglomerular apparatus (JGA)

○ Granular cells

○ Function : Sensing of afferent arteriole pressure, secretion of renin.


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Figure. 7. Juxtaglomerular Apparatus


② 1st. Granular cells in the afferent arteriole secrete renin.

○ Renin is a type of protein-degrading enzyme.

③ 2nd. Renin activates angiotensinogen secreted in the liver to angiotensin I.

④ 3rd. Angiotensin I is converted to angiotensin II by angiotensin-converting enzyme.

○ Angiotensin-converting enzyme (ACE) : Located in the lungs. Chymase.

○ ACE2 : Prolyl carboxypeptidase.

⑤ 4th. Angiotensin II has the following functions through signal transduction via angiotensin II receptors.

○ Angiotensin II receptors : AT1 (Angiotensin II type 1 receptor), etc.

Function 1. Stimulation of adrenal cortex → Secretion of aldosterone → Reabsorption of Na+ in the blood → Increase in blood pressure.

Function 2. Constriction of afferent arterioles and efferent arterioles → Increase in blood pressure.

Function 3. Stimulation of ADH secretion in the posterior pituitary → Promotion of water reabsorption in the kidneys → Increase in blood pressure.

Function 4. Promotion of Na+ reabsorption in the kidneys.

Function 5. Inhibition of bradykinin, a vasodilator.

⑥ 5th. Stimulated adrenal cortex secretes aldosterone (a mineralocorticoid).

⑦ 6th. Aldosterone promotes Na+ reabsorption in the distal convoluted tubule and the collecting duct.

○ Na+ reabsorption : Involves the Na+/K+ pump, leading to increased secretion of K+ (Na+ : K+ = 3 : 2).

○ Na+ reabsorption (active transport) Reabsorption of Cl- (passive transport) and water (passive transport) for osmotic balance.

⑧ 7th. Effect of increased blood volume on blood pressure.

⑨ 8th. Renin negative feedback

○ 8th - 1st. ATP is secreted into the blood vessels through the Na+/K+ pump.

○ 8th - 2nd. Simultaneously, ATP is used in the Na+/K+ pump, generating ADP.

○ 8th - 3rd. ADP generates adenosine.

○ 8th - 4th. Adenosine is secreted into the blood vessels.

○ 8th - 5th. Both ATP and adenosine increase Ca2+ in vascular smooth muscle cells.

○ 8th - 6th. Increased Ca2+ moves to renin-secreting cells, inhibiting renin secretion.


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Figure. 8. Renin Negative Feedback


⑨ Types of angiotensin

○ Angiotensinogen : N-terminal - Asp - Arg - Val - Tyr - Ile - His - Pro - Phe - His - Leu - Leu - Val - Tyr - Ser - R

○ Angiotensin I : Asp - Arg - Val - Tyr - Ile - His - Pro - Phe - His - Leu

○ Angiotensin II (1-8) : Asp - Arg - Val - Tyr - Ile - His - Pro - Phe

○ Angiotensin III (2-8) : Arg - Val - Tyr - Ile - His - Pro - Phe

○ Angiotensin IV (3-8) : Val - Tyr - Ile - His - Pro - Phe

○ Angiotensin V ( (1-7) : Asp - Arg - Val - Tyr - Ile - His - Pro

⑩ Plasma sodium concentration homeostasis is primarily influenced by ADH rather than the RAAS.

⑪ ACE inhibition

○ Research to find substances inhibiting ACE is actively underway to prevent hypertension.

Measurement method 1. Measuring the conversion of HHL (hippuryl-histidyl-leucine) to HA (hippuric acid) by ACE.

○ Measuring HA using UV-vis spectroscopy or HPLC.

○ Disadvantage : Organic solvents like ethyl acetate are used in the HA extraction process, reducing accuracy.

Measurement method 2. ACE kit-WST : Measuring the conversion of 3HB-GGG (3-hydroxybutyryl-gly-gly-gly) to 3HB (3-hydroxybutyric acid) by ACE.


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Figure. 9. ACE kit-WST


⑶ Atrial Natriuretic Peptide (ANP)

① Secreted from atrial pressure receptors in the left atrium : Secretion increases when atrial receptors are stretched more than normal.

② Inhibits RAAS, promotes ADH antagonism.

③ Promotes excretion of salt (sodium, etc.) and water in the kidneys → Decreases blood pressure → Reduces body weight.

④ Prevents strain on the heart due to excessive fluid.


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Figure. 10. Regulation of Blood Filtration


⑷ Autonomic Nervous System

① Sympathetic Nervous System : Relaxation of bladder smooth muscles (sensation of needing to urinate), contraction of urethral smooth muscles (prevents urine release).

② Parasympathetic Nervous System : Contraction of bladder smooth muscles, relaxation of urethral smooth muscles.

③ Somatic neurons (motor neurons) → Contraction of detrusor muscle in the bladder.



7. Renal Disorders

⑴ Syndrome of Inappropriate Antidiuretic Hormone Secretion (SIADH)

⑵ Diabetes Insipidus

① Central Diabetes Insipidus: Issues with the hypothalamus or pituitary gland → Normal response to ADH administration

② Nephrogenic Diabetes Insipidus: Issues with ADH receptors

⑶ Acute Kidney Injury (AKI)

① COVID-19 infection can also cause acute kidney injury

② Treatment: AMF (amifostine)

⑷ Chronic Kidney Disease (CKD)


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Table. 4. Classification of CKD


⑸ Hypercalcemia

① Acidosis, Alkalosis

② Hyperaldosteronism: Metabolic alkalosis

③ Hypoaldosteronism: Metabolic acidosis

④ Hypokalemia → Decreased H+

⑹ Renal Transplantation

① In renal transplantation, the kidney must be removed, and a new kidney must be transplanted within 48 hours.



8. Sweat

⑴ Functions of Sweat

① Temperature regulation: Utilizes the evaporation of sweat to lower elevated body temperature

② Edema prevention: Intense exercise → Increased systolic blood pressure → Increased capillary permeability → Edema → Prevention of edema through loss of moisture via sweat

⑵ Sweat glands secrete Ach under the influence of postganglionic nerves.



Input: 2015.07.24 11:25

Modified: 2019.12.19 00:07

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