Chapter 41

Chemical Signals in Animals

•授課教師：臺北醫學大學 彭志維老師
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•電子郵件：cwpeng@tmu.edu.tw

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 Overview: The Body’s Long-Distance Regulators
• Animal hormones are chemical signals that are secreted into
the circulatory system and communicate regulatory messages
within the body
• Hormones reach all parts of the body, but only target cells
have receptors for that hormone
• Insect metamorphosis (變態) is regulated by hormones

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 • Two systems coordinate communication
throughout the body: the endocrine system and
the nervous system
• The endocrine system(內分泌系統) secretes
hormones that coordinate slower but longer-
acting responses including reproduction,
development, energy metabolism, growth, and
behavior
• The nervous system (神經系統)conveys high-
speed electrical signals along specialized cells
called neurons; these signals regulate other cells
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 Concept 41.1: Hormones and other signaling
molecules bind to target receptors, triggering
specific response pathways
• Endocrine signaling is just one of several ways
information is transmitted between animal cells
• The ways that signal are transmitted between animal
cells are classified by two criteria
– The type of secreting cell
– The route taken by the signal in reaching its target
• Hormones secreted into extracellular fluids by endocrine
cells reach their targets via the bloodstream
• Endocrine signaling maintains homeostasis, mediates
responses to stimuli, regulates growth and development 4
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 Paracrine(旁泌) and Autocrine (自泌)Signaling

• Local regulators(局部傳訊) are molecules

that act over short distances, reaching
target cells solely by diffusion
– In paracrine signaling(旁泌傳訊), the target
cells lie near the secreting cells
– In autocrine signaling(自泌傳訊), the target
cell is also the secreting cell

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Figure 45.2

Blood
vessel Response

內分泌傳訊 (a) Endocrine signaling

Response

旁泌傳訊 (b) Paracrine signaling

Response

自泌傳訊 (c) Autocrine signaling

Synapse

Neuron

Response

突觸傳訊 (d) Synaptic signaling

Neurosecretory
cell

Blood
vessel Response

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神經內分泌傳訊 (e) Neuroendocrine signaling
 Synaptic and Neuroendocrine Signaling
• In synaptic signaling(突觸傳訊), neurons form
specialized junctions with target cells, called
synapses(突觸)
• At synapses, neurons secrete molecules called
neurotransmitters(神經傳遞物質) that diffuse
short distances and bind to receptors on target
cells
• In neuroendocrine signaling(神經內分泌傳訊),
specialized neurosecretory cells secrete molecules
called neurohormones(神經賀爾蒙) that travel to
target cells via the bloodstream
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 Signaling by Pheromones (費洛蒙)
• Members of the same animal species sometimes
communicate with pheromones (費洛蒙),
chemicals that are released into the environment
• Pheromones serve many functions, including
marking trails leading to food, a wide range of
functions that include defining territories, warning
of predators, and attracting potential mates

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 Endocrine Tissues and Organs
• In some tissues, endocrine cells are grouped
together in ductless organs called endocrine
glands(內分泌腺)
• Endocrine glands secrete hormones directly into
surrounding fluid
• These contrast with exocrine glands (外分泌腺),
which have ducts and which secrete substances
onto body surfaces or into cavities

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Figure 45.4
Major endocrine glands:
下視丘 Hypothalamus
松果體 Pineal gland
腦下垂體 Pituitary gland Organs containing
endocrine cells:
甲狀腺 Thyroid gland
Thymus 胸腺
副甲狀腺 Parathyroid glands
(behind thyroid) Heart 心臟
Liver
Adrenal glands
腎上腺 (atop kidneys)
Stomach

胰臟 Kidneys
Pancreas
(胰島)
Small
卵巢 Ovaries (female) intestine

睪丸 Testes (male)

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 Chemical Classes of Hormones
• Three major classes of molecules function as
hormones in vertebrates
– Polypeptides (多肽類激素；proteins and peptides)
– Amines derived from amino acids(胺類激素)
– Steroid hormones (類固醇激素)
• Lipid-soluble hormones (steroid hormones) pass
easily through cell membranes, while water-soluble
hormones (polypeptides and amines) do not
• The solubility of a hormone correlates with the
location of receptors inside or on the surface of
target cells
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Figure 45.5
Water-soluble (hydrophilic) Lipid-soluble (hydrophobic)

多肽類激素 Polypeptides Steroids

類固醇激素

0.8 nm
Insulin Cortisol

Amines
胺類激素

Epinephrine Thyroxine 12
 Cellular Response Pathways
• Water and lipid soluble hormones differ in their
paths through a body
• Water-soluble hormones are secreted by
exocytosis(胞吐作用), travel freely in the
bloodstream, and bind to cell-surface receptors
• Lipid-soluble hormones diffuse across cell
membranes, travel in the bloodstream bound to
transport proteins(載體蛋白), and diffuse through
the membrane of target cells

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Figure 45.6-2
SECRETORY
CELL

Water- Lipid-
soluble soluble
hormone hormone

VIA
BLOOD
Transport
Signal receptor protein

TARGET OR
CELL Signal
receptor

Cytoplasmic
response Gene
regulation
Cytoplasmic
response Gene
regulation

NUCLEUS
(a) (b) 14
 Pathway for Water-Soluble Hormones
• Binding of a hormone to its receptor initiates a
signal transduction pathway leading to responses
in the cytoplasm, enzyme activation, or a change in
gene expression
• The hormone epinephrine (腎上腺)has multiple
effects in mediating the body’s response to short-
term stress
• Epinephrine binds to receptors on the plasma
membrane of liver cells
• This triggers the release of messenger molecules
that activate enzymes and result in the release of
glucose into the bloodstream
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Figure 45.7-2

Epinephrine EXTRACELLULAR
FLUID
G protein Adenylyl
cyclase

G protein-coupled GTP
receptor
ATP

cAMP Second
messenger

Inhibition of Protein
glycogen synthesis kinase A

Promotion of
glycogen breakdown

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 Pathway for Lipid-Soluble Hormones
• The response to a lipid-soluble hormone is
usually a change in gene expression
• Steroids, thyroid (甲狀腺)hormones, and the
hormonal form of vitamin D enter target cells and
bind to protein receptors in the cytoplasm or
nucleus
• Protein-receptor complexes then act as
transcription factors in the nucleus, regulating
transcription of specific genes

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Figure 45.8-2
Hormone EXTRACELLULAR
(estradiol) FLUID

Estradiol
(estrogen)
receptor Plasma
membrane

Hormone-receptor
complex
NUCLEUS

CYTOPLASM

DNA
Vitellogenin
mRNA
for vitellogenin

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 Multiple Effects of Hormones
• The same hormone may have different effects on
target cells that have
– Different receptors for the hormone
– Different signal transduction pathways
• For example, epinephrine simultaneously
triggers glycogen breakdown in the liver,
decreased blood flow to the digestive tract,
and increased blood flow to major skeletal
muscles.

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肝細胞 骨骼肌血管 腸道肌血管

放鬆 收縮

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 Signaling by Local Regulators
• Local regulators are secreted molecules that link
neighbouring cells or directly regulate the secreting
cell
• Types of local regulators
– Cytokines (細胞素) and growth factors(生長因子)
– Nitric oxide (NO；一氧化氮)
– Prostaglandins (前列腺素)
• In the immune system, prostaglandins promote fever
and inflammation and intensify the sensation of pain
• Prostaglandins help regulate aggregation of platelets,
an early step in formation of blood clots
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 Coordination of Neuroendocrine and
Endocrine Signaling
• The endocrine and nervous systems generally
act coordinately to control reproduction and
development
• For example, in larvae (幼蟲)of butterflies and
moths, the signals that direct molting(蛻皮)
originate in the brain

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 • In insects, molting and development are controlled
by a combination of hormones
– A brain hormone (PTTH) stimulates release of
ecdysteroid(蛻皮素) from the prothoracic glands
(前胸腺)
– Juvenile hormone(保幼激素) promotes retention of
larval characteristics
– Ecdysone (蛻皮素) promotes molting (in the
presence of juvenile hormone) and development
(in the absence of juvenile hormone) of adult
characteristics

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Figure 45.10-3

昆蟲的發育，幼蟲期
Brain 神經分泌細胞
Neurosecretory cells
經過一次又一次的蛻
皮(舊的外骨骼)，發 Corpora cardiaca
育成蛹，經過變態的 Corpora allata 咽喉側線
PTTH
過程發育成成體型態 腦部神經分泌細胞合成腦激素
Prothoracic 儲存於心腺中，釋出後傳送訊
gland 息給前胸腺合成蛻皮素，蛻皮
前胸腺 素的分泌為間歇性的，每一次
Juvenile
hormone (JH) 釋出就發生一次蛻皮，保幼激
保幼激素 素則由咽喉側腺分泌，此激素
Low
Ecdysteroid JH 可以刺激幼蟲長大同時抑制變
態反應發生
蛻皮素

EARLY
LATER
LARVA
LARVA
PUPA 蛹 ADULT
幼蟲早期 幼蟲晚期 當保幼激素濃度下降，則蛻皮素刺
激幼蟲發育成蛹 變態反應 24
 Concept 41.2: Simple Hormone Pathways
• Two basic types: simple endocrine and simple
neuroendocrine pathways
• In a simple endocrine pathway, endocrine cells
respond directly to an internal or environmental
stimulus by secreting a particular hormone.
• For example, the release of acidic contents of the
stomach into the duodenum stimulates endocrine
cells there to secrete secretin (胰泌素)
• This causes target cells in the pancreas (胰臟), a
gland behind the stomach, to raise the pH in the
duodenum
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Figure 41.10
Simple endocrine pathway Example: secretin signaling

STIMULUS Low pH in
duodenum
十二指腸
Endocrine S cells of duodenum
cell
Negative feedback

Hormone Secretin (•)

胰泌素

負回饋

Target Pancreatic cells

cells

RESPONSE Bicarbonate release

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 • In a simple neuroendocrine pathway, the stimulus
is received by a sensory neuron, which stimulates
a neurosecretory cell
• The neurosecretory cell secretes a
neurohormone, which enters the bloodstream
and travels to target cells

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Figure 41.11
Simple neuroendocrine pathway Example: oxytocin signaling

STIMULUS Suckling

Sensory neuron

Hypothalamus/
posterior pituitary
Positive feedback

Neurosecretory
cell Oxytocin (•)
正回饋 Neurohormone
(催產素)

Target Smooth muscle in

cells mammary glands

RESPONSE Milk release 28

 Feedback Regulation

• A negative feedback loop inhibits a response by

reducing the initial stimulus, thus preventing
excessive pathway activity
• Positive feedback reinforces a stimulus to
produce an even greater response
• For example, in mammals oxytocin (催產素)
causes the release of milk, causing greater
suckling by offspring, which stimulates the
release of more oxytocin

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 Insulin and Glucagon: Control of Blood
Glucose
• Insulin (胰島素,decreases blood glucose) and
glucagon (升糖素,increases blood glucose) are
antagonistic hormones that help maintain glucose
homeostasis
• The pancreas has clusters of endocrine cells
called pancreatic islets (胰島) with alpha cells
that produce glucagon and beta cells that
produce insulin

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Figure 45.13
Body cells Insulin
take up more Beta cells of
glucose. pancreas
release insulin
into the blood. 胰島素
Liver takes
up glucose
and stores it STIMULUS:
as glycogen. Blood glucose level rises
Blood glucose
level declines. (for instance, after eating a
carbohydrate-rich meal).

Homeostasis:
Blood glucose level
(70–110 mg/m100mL)

STIMULUS:
Blood glucose Blood glucose level
level rises. falls (for instance, after
skipping a meal).

Liver breaks
down glycogen Alpha cells of pancreas
release glucagon into
and releases
glucose into the blood. 升糖素
the blood. Glucagon 31
 Target Tissues for Insulin and Glucagon
• Insulin (胰島素) reduces blood glucose levels by
– Promoting the cellular uptake of glucose
– Slowing glycogen breakdown in the liver
– Promoting fat storage, not breakdown
• Glucagon (升糖素) increases blood glucose
levels by
– Stimulating conversion of glycogen to glucose
in the liver
– Stimulating breakdown of fat and protein into
glucose
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 Diabetes Mellitus
• Diabetes mellitus (糖尿病) is perhaps the best-
known endocrine disorder
• It is caused by a deficiency of insulin or a
decreased response to insulin in target tissues
• It is marked by elevated blood glucose levels
• Type I diabetes mellitus (insulin-dependent) is an
autoimmune disorder in which the immune
system destroys pancreatic beta cells
• Type II diabetes mellitus (non-insulin-dependent)
involves that target cells fail to take up gulocose
due to change in insulin receptors
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 41.3: The hypothalamus(下視丘) and pituitary
(腦下腺) are central to endocrine regulation
• The hypothalamus receives information from the
nervous system and initiates responses through the
endocrine system
• Attached to the hypothalamus is the pituitary
gland composed of the posterior pituitary and
anterior pituitary
• The posterior pituitary stores and secretes
hormones that are made in the hypothalamus
• The anterior pituitary makes and releases
hormones under regulation of the hypothalamus
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Figure 45.14
Cerebrum
Pineal
gland Thalamus

Hypothalamus
Cerebellum
Pituitary
Spinal cord gland

Hypothalamus
Posterior 下視丘
pituitary
垂體後葉
Anterior
pituitary
垂體前葉 35
 Posterior Pituitary Hormones
• The two hormones released from the posterior
pituitary act directly on nonendocrine tissues
– Oxytocin(催產素) regulates milk secretion by the
mammary glands
– Antidiuretic hormone (抗利尿素；ADH)
regulates physiology and behavior

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Figure 45.15
Hypothalamus

Neurosecretory
cells of the
hypothalamus

Neurohormone Axons

Posterior
腦下腺後葉 pituitary
(神經性腦下腺) Anterior
pituitary

HORMONE ADH Oxytocin

抗利尿素 催產素
TARGET Kidney Mammary glands,
tubules uterine muscles 37
 Anterior Pituitary Hormones
• Hormone production in the anterior pituitary is
controlled by releasing and inhibiting hormones
from the hypothalamus
• For example, prolactin-releasing hormone (泌乳
素釋放激素) from the hypothalamus stimulates
the anterior pituitary to secrete prolactin (泌乳素；
PRL), which has a role in milk production

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促進性激素

非促進性激素

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Table 45.1a

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Table 45.1b

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Thyroid Regulation: A Hormone Cascade
Pathway
• A hormone can stimulate the release of a series
of other hormones, the last of which activates a
nonendocrine target cell; this is called a
hormone cascade pathway
• The release of thyroid hormone results from a
hormone cascade pathway involving the
hypothalamus, anterior pituitary, and thyroid
gland
• Hormone cascade pathways typically involve
negative feedback 42
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 Figure 41.16
STIMULUS 1 Thyroid hormone
levels drop. 甲狀腺素下降
Sensory
neuron
2 The hypothalamus secretes
Hypothalamus TRH ●into the blood. Portal 甲狀腺刺激素
vessels carry TRH to anterior
釋放激素

Negative feedback
Neuro- pituitary.
secretory
cell
TRH

3 TRH causes anterior pituitary

to secrete TSH▲ . 甲狀腺刺激素
負回饋 TSH
Anterior
pituitary
Circulation
throughout
body via blood
Thyroid
gland
4 TSH stimulates endocrine
cells in thyroid gland to 甲狀腺素
secrete T3 and T4 ■.
Thyroid
hormone
6 Thyroid hormone blocks TRH
release and TSH release Circulation
preventing overproduction throughout
of thyroid hormone. body via blood

RESPONSE
5 Thyroid hormone levels
return to normal range.
甲狀腺素濃度正常
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Disorders of Thyroid Function and Regulation
• Hypothyroidism(甲狀腺機能不足), too little thyroid
function, can produce symptoms such as
– Weight gain, lethargy (昏睡), cold intolerance
• Hyperthyroidism (甲狀腺機能亢進), excessive
production of thyroid hormone, can lead to
– High temperature, sweating, weight loss,
irritability and high blood pressure
• Malnutrition can alter thyroid function
• Insufficient dietary iodine leads to an enlarged
thyroid gland, called a goiter(甲狀腺腫大)
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 • Graves disease(格雷夫氏症), a form of hyperthyroidism
caused by autoimmunity, is typified by protruding eyes
• Thyroid hormone refers to a pair of hormones
– Triiodothyronin (T3), with three iodine atoms
– Thyroxine (T4) with four iodine atoms

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 Tropic (促進性) and Nontropic Hormones
• A tropic hormone regulates the function of endocrine
cells or glands. Three primarily tropic hormones are
– Follicle-stimulating hormone (FSH;濾泡細胞刺激素)
– Luteinizing hormone (LH;黃體生成素)
– Adrenocorticotropic hormone (ACTH;腎上腺皮質刺
激素)
• FSH and LH are also called gonadotropins(促性腺激素
). Their hypothalamic regulator is called gonadotropin-
releasing hormone (GnRH;促性腺激素釋放素).
• ACTH stimulates the production and secretion of
steroid hormones by the adrenal cortex.
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 • Growth hormone (GH;生長激素) is secreted by
the anterior pituitary gland and has tropic and
nontropic actions
• It promotes growth directly and has diverse
metabolic effects
• It stimulates production of growth factors
• An excess of GH can cause gigantism(巨人症),
while a lack of GH can cause dwarfism(侏儒症)

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Concept 41.4: Endocrine glands respond to
diverse stimuli in regulating homeostasis,
development, and behavior
• Endocrine signaling regulates homeostasis, development,
and behavior
Parathyroid Hormone and Vitamin D: Control
of Blood Calcium
• Two antagonistic hormones regulate the homeostasis of
calcium (Ca2+) in the blood of mammals
– Parathyroid hormone (PTH;副甲狀腺素) is released
by the parathyroid glands
– Calcitonin (降鈣素) is released by the thyroid gland

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 • PTH (副甲狀腺素)increases the level of blood Ca2+
– It releases Ca2+ from bone and stimulates
reabsorption of Ca2+ in the kidneys
– It also has an indirect effect, stimulating the kidneys
to activate vitamin D, which promotes intestinal
uptake of Ca2+ from food
• Calcitonin(降鈣素) decreases the level of blood Ca2+
– It stimulates Ca2+ deposition in bones and secretion
by kidneys

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Figure 41.19

NORMAL BLOOD
Ca2+ LEVEL
(about 10 mg/100 mL)
Blood Ca2+
level rises.

Blood Ca2+ level falls.

Active vitamin D
increases Ca2+.

PTH stimulates Ca2+

uptake and promotes
activation of vitamin D.

Parathyroid
glands
PTH release PTH.
PTH stimulates
Ca2+ release. 副甲狀腺素 50
 Adrenal Hormones: Response to Stress
• Each adrenal gland (腎上腺) actually consists of two
glands: adrenal medulla (inner portion;腎上腺髓質) and
adrenal cortex (outer portion;腎上腺皮質)

Catecholamines (兒茶酚胺) from the Adrenal Medulla

• The adrenal medulla secretes epinephrine(adrenaline;腎
上腺素) and norepinephrine (noradrenaline;正腎上腺素)
• These hormones are members of a class of compounds
called catecholamines
• They are secreted in response to stress-activated
impulses from the nervous system
• They mediate various fight-or-flight responses

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 • Epinephrine and norepinephrine
– Trigger the release of glucose and fatty acids into
the blood
– Increase oxygen delivery to body cells
– Direct blood toward heart, brain, and skeletal
muscles, and away from skin, digestive system,
and kidneys
• The release of epinephrine and norepinephrine
occurs in response to involuntary nerve signals

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Figure 41.20a
(a) Short-term stress response and the adrenal medulla
Stress

Spinal cord Nerve

Hypo-
(cross section) signals thalamus

Nerve
cell

Nerve cell
Adrenal medulla
secretes epinephrine
and norepinephrine.

Effects of epinephrine and norepinephrine: Adrenal

gland
• Glycogen broken down to glucose;
increased blood glucose Kidney
• Increased blood pressure
• Increased breathing rate
• Increased metabolic rate
• Change in blood flow patterns, leading to
increased alertness and decreased digestive,
excretory, and reproductive system activity 53
Steroid(類固醇) Hormones from the Adrenal Cortex
• The adrenal cortex releases a family of steroids called
corticosteroids(皮質類固醇) in response to stress
• These hormones are triggered by a hormone cascade
pathway via the hypothalamus and anterior pituitary
(ACTH；腎上腺皮質刺激素)
• Humans produce two types of corticosteroids:
glucocorticoids (糖皮質素) and mineralocorticoids (鹽皮質
素)
• Glucocorticoids, such as cortisol(皮質醇), influence
glucose metabolism and the immune system
• Mineralocorticoids, such as aldosterone(醛固酮), affect
salt and water balance
• The adrenal cortex also produces small amounts of
steroid hormones that function as sex hormones 54
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Figure 41.20b

(b) Long-term stress response and the adrenal cortex

Stress
Hypothalamus
Releasing
hormone

Anterior pituitary
Blood vessel
ACTH Effects of Effects of
mineralocorticoids: glucocorticoids:

• Retention of sodium • Proteins and fats broken

Adrenal ions and water by down and converted to
gland kidneys glucose, leading to
Adrenal cortex increased blood glucose
secretes mineralo- • Increased blood
corticoids and volume and blood • Partial suppression of
glucocorticoids. pressure immune system
Kidney

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 Gonadal Sex Hormones
• The gonads, testes and ovaries, produce most of
the sex hormones: androgens(雄性賀爾蒙),
estrogens(動情素), and progestins(黃體素)
• All three sex hormones are found in both males and
females, but in significantly different proportions
• The testes primarily synthesize androgens, mainly
testosterone (睪固酮), which stimulate
development and maintenance of the male
reproductive system
• Testosterone causes an increase in muscle and
bone mass and is often taken as a supplement to
cause muscle growth, which carries health risks
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 • Estrogens (動情素), most importantly estradiol(雌
二醇), are responsible for maintenance of the
female reproductive system and the development
of female secondary sex characteristics
• In mammals, progestins(黃體素), which include
progesterone(黃體酮), are primarily involved in
preparing and maintaining the uterus
• Synthesis of the sex hormones is controlled by FSH
and LH from the anterior pituitary
• FSH and LH production is controlled by a releasing
hormone from the hypothalamus, GnRH
(gonadotropin-releasing hormone).促性腺激素釋放素57
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 Endocrine Disruptors(內分泌干擾物)
• Between 1938 and 1971 some pregnant women
at risk for complications were prescribed a
synthetic estrogen called diethylstilbestrol (DES;
乙烯雌酚)
• Daughters of women treated with DES are at
higher risk for reproductive abnormalities,
including miscarriage, structural changes, and
cervical and vaginal cancers
• DES is an endocrine disruptor, a molecule that
interrupts the normal function of a hormone
pathway, in this case, that of estrogen
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