BIO270 L10

BIO270: Lecture Topic Overview

Lecture 1: Basic Physiological Principles
Lecture 2: Biochemical Basis of Physiology: Part I
Lecture 3: Biochemical Basis of Physiology: Part II
Lecture 4: Cell Signaling
Lecture 5: Signal Transduction/Endocrine Regulation
Midterm
Lecture 6: Reproduction
Lecture 7: Movement and Muscle Physiology: Part I
Lecture 8: Muscle Physiology: Part II
Lecture 9: Osmoregulation: Ion and Water Balance
Chapter 10: pages 470-514
Lecture 10: Digestion
Chapter 11: pages 528-538, 543-565 (not box 11.3)
Lecture 11: Thermal Physiology
Final Exam
1
 BIO270 L10
BIO270 L10 Overview

Feeding and Digestion

1.  Overview
2.  Diet and energy
3.  Nutrient breakdown and transport
4.  Digestive systems
5.  Regulation of feeding and digestion

2
 BIO270 L10
Overview of digestion
Assimilation – processes of nutrient acquisition, digestion and absorption

§  Most chemical breakdown occurs

externally, within the lumen of the
GI tract
§  Complex organ, many cell types:
§  Secretory
§  Absorptive
§  Muscle cells
§  Neurons

§  Assimilated nutrients are broken

down for energy or used as
building blocks

§  Egestion: the elimination of

undigested food
3
 BIO270 L10
BIO270 L10 Overview

Feeding and Digestion

1.  Overview
2.  Diet and energy
3.  Nutrient breakdown and transport
4.  Digestive systems
5.  Regulation of feeding and digestion

4
 BIO270 L10
Diets Provide Energy
§  Energy content of diet must match the metabolic demands of the
animal
§  Short term imbalances buffered by fuel storage depots
§  Caloric equivalent
§  Energy content of a gram of a specific macromolecule
§  Protein and carbohydrates = 4 kcal/gm
§  Fat = 9 kcal/gm
§  Gross energy – total chemical energy
§  Measured by calorimetry
§  Food is burned, heat released is measured
§  Net energy - food energy available to animals through
assimilation

Nutrients are the external molecules that allow an

animal to maintain and build cells.
5
 BIO270 L10
Diets Provide Energy

Some food is indigestible or unmetabolizable

(Heat Increment)

Energy lost through the increase in metabolic

rate during digestion.
Important source of thermal energy.
6
 BIO270 L10
Vitamins
§  Group of unrelated molecules with diverse functions
§  Many participate in catalysis as cofactors for enzymes
§  Usually categorized based on solubility
§  Fat-soluble – vitamins A, D, E, K (synthesized by body)
§  Stored in fat; can have toxicity
§  Water soluble – vitamins B, C
§  Excess excreted in urine

§  Ship’s surgeon James Lind in 1747

identified a quality in fruit

§  Obtained in diet or from bacteria living in the GI tract

7
 BIO270 L10
Minerals
§  Metallic elements that participate in protein structure
§  Calcium
§  Phosphorus
§  Iron – eg. structure of hemoglobin protein
§  Copper – eg. electron and oxygen transport
§  Zinc – binds many proteins, ligands in metabolism,
signalling etc.
§  Most are absorbed along the GI tract by specific
transporters residing in the GI tract

8
 BIO270 L10
Amino Acids
§  Animals use 20 amino acids to build proteins
§  Most can be produced by the animal – de novo synthesis
§  Eight essential amino acids must be obtained in the diet
§  Depends on species (eg. dogs, cats have additional requirements)
§  Diets deficient in any essential amino acid lead to developmental
defects and slow growth

§  Protein quality

§  The amino acid profile of dietary protein
§  Animal tissue provides higher protein quality than plant tissue
§  Some plants lack specific essential amino acids
§  Herbivores avoid amino acid deficiencies by eating a variety of plants

9
 BIO270 L10
Fatty acids
§  Animals can make almost all fatty acids from acetyl
CoA
§  Animals cannot produce sufficient amounts of
omega-3 (ω3) or omega-6 (ω6) fatty acids

§  Omega-3 (ω3) fatty acids must be ingested as linolenic (18:3

ω3) acid
§  Found in cold-water fish
§  Omega-6 (ω6) fatty acids must be ingested as linoleic (18:2
ω6)
§  Found in plant seeds, oils, nuts etc.

10
 BIO270 L10
Digestive Enzymes
Allow animals to convert complex macromolecules to
forms that can be absorbed and processed
§  Lipases
§  Break down triglycerides and phospholipids into fatty acids
§  Proteases
§  Break down proteins to shorter polypeptides
§  Amylases
§  Break down polysaccharides into oligosaccharides
§  Nucleases
§  Break down DNA into nucleotides

Most digestion takes place extracellularly

§  For example, in the lumen of the GI tract
11
 BIO270 L10
BIO270 L10 Overview

Feeding and Digestion

1.  Overview
2.  Diet and energy
3.  Transport and nutrient breakdown
4.  Digestive systems
5.  Regulation of feeding and digestion

12
 BIO270 L10
Nutrient transport across plasma membranes

§  Some nutrients are transported by protein carriers

§  Polar molecules require specific protein transporters eg.
monosaccharides and amino acids
§  Transport down a concentration gradient
§  Facilitated diffusion
§  Eg. Glucose transporters (GLUT proteins)
§  Transport against a concentration gradient
§  Active transport
§  Via Na+-dependent cotransporters

§  Some nutrients are transported in vesicles

§  Pinocytosis (in solution nutrients)/phagocytosis (particulate
nutrients) combined with exocytosis

13
 BIO270 L10
Carbohydrate breakdown

§  Main types consumed by animals

§  Polysaccharides
§  Glycogen, starch, cellulose, chitin
§  Disaccharides
§  Sucrose, lactose, maltose

§  Polysaccharides and disaccharides are broken down to

monosaccharides
§  Glucose, fructose, galactose

14
 BIO270 L10
Carbohydrate breakdown and absorption

Monosaccharides are
absorbed by
specialized epithelial
cells in the small
intestine (enterocytes)
15
 BIO270 L10
Carbohydrate absorption by enterocytes
§  Combination of active transport (SGLT-1) and
facilitated diffusion (GLUT proteins)

Preformed GLUT2
are also transported
to apical membrane

16
 BIO270 L10
Lactose Intolerance
3 types:
Primary: amount of lactase reduced into adulthood (genetic)
Secondary (acquired): result of inflammation (i.e. Crohn’s, gastroenteritis, parasites etc.)
Congenital: rare autosomal recessive genetic disorder

Retrieved from
http://www.avonmorelactosefree.ie/lactose-
intolerance/what-is-lactose-intolerance.html,
15,11,2013 17
 BIO270 L10
Protein breakdown

§  Extracellular hydrolysis,

starting in the stomach, to
dipeptides and amino acids
§  Amino acids are transported
into epithelial cells by amino
acid-Na+ cotransporters
§  Some proteins are also carried
into cells intact via transcytosis
§  E.g. Infant mammalian
antibodies in collostrum
§  Transferring immunoprotection

18
 BIO270 L10
Lipids
§  Digestion and import of lipids is
complicated by their hydrophobicity
§  GI tract secretes bile that emulsifies lipids
into small droplets (micelles)
§  Accessible to pancreatic lipase
§  Dietary fats are broken down into fatty acids
and monoglycerides Short chain

§  Lipids diffuse across cell membrane into

enterocyte
§  Transport of lipids depend on physical
properties
§  Short chain fatty acids and glycerol in blood
§  Sufficiently polar
§  Triglycerides in the lymph as chylomicrons
§  Processed by lipoprotein lipase (glycerol and
FAs)
§  Taken up by peripheral tissues; repackaged
by liver 19
 BIO270 L10
BIO270 L10 Overview

Feeding and Digestion

1.  Overview
2.  Diet and energy
3.  Transport and nutrient breakdown
4.  Digestive systems
5.  Regulation of feeding and digestion

20
 BIO270 L10
Digestive systems
§  Evolutionary history shows increasing anatomical
and functional specialization
§  Two-way gut
§  Simple internal sac
§  Isolate in a controlled environment
§  Sac may have diverticula to increase surface area
§  Food enters and wastes leave via the same opening
§  Sponges, cnidarians, flatworms

21
 BIO270 L10
Digestive systems
§  One-way gut
(gastrointestinal tract)
§  Specialized regions
§  Mouth, pharynx, esophagus
§  Mechanical breakdown of food
§  Stomach
§  Acidic compartment
§  Upper or small intestines
§  Digestion and absorption
§  Lower or large intestines
§  Absorption of water
§  Anus
§  Release of indigestible material
22
 BIO270 L10
Surface area
§  In most animals, nutrients are hydrolyzed in the
lumen of the GI tract
§  Nutrients are taken up by cells lining the gut
§  Nutrient uptake is improved by increasing surface
area in two ways
1.  Increasing gut length
2.  Increasing surface undulations
§  Circular folds
§  Villi
§  Microvilli

23
 BIO270 L10
Specialized compartments
§  Specialized compartments increase efficiency of digestion
§  More developed in animals with one-way gut
§  Compartments have functional specializations
§  Altering the pH
§  Secretions: enzymes, mucus
§  Region-specific types of secretory and absorptive cells
§  Muscular valves (sphincters) control passage of food from one
compartment to the next
§  Complexity of gut morphology varies across taxa
§  Reflects complexity of the diet, ease of digestion, and life history
§  Also within taxa

24
 BIO270 L10
Variations in vertebrate gut morphology

Ceca: branches of digestive tract that house bacteria-enterosymbionts

Crop: outpouching of esophagus for storing partially digested food
25
 BIO270 L10
Symbiotic organisms
§  Digestion in many animals benefits from the assistance of
symbiotic organisms
§  For example, bacteria, fungi, and photosynthetic organisms

§  Three main types of symbionts participate in digestion

§  Enterosymbionts
§  Live within the lumen of the GI tract
§  Often in an enlarged region called the cecum

§  Exosymbionts
§  Actively cultivated outside the body

§  Endosymbionts
§  Grow in interstitial spaces or (less often)
within host cells

26
 BIO270 L10
Ruminants
§  Many mammals possess modifications that improve the digestion of plant
material. E.g. Ruminants (grazing animals: cows, sheep, giraffe, deer) possess a
digastric stomach:
1.  Rumen and reticulum – fermentative bacteria housed here
2.  Omasum (water absorption) and abomasum (digestive enzymes)

27
 BIO270 L10
Ruminants digest cellulose

28
 BIO270 L10
Salivary glands
§  Collection of several exocrine glands
§  Ducts open into mouth
§  Multicellular
§  Mucus-secreting cells
§  Serous cells – secrete enzymes
§  Saliva
§  Lubricates food
§  Dissolves food so nutrients can bind to gustatory receptors
§  Cleanses the mouth with antimicrobial properties
§  Contain enzymes that initiate digestion
§  Salivation is controlled by nerve signals
§  Parasympathetic nerves stimulate salivation
§  Sympathetic nerves inhibit salivation
29
 BIO270 L10
Stomach
§  Surface is composed of columnar epithelial cells
joined by tight junctions
§  Prevent leakage across epithelium
§  Surface mucosa contains gastric pits lined by
4 cell types:
1.  Mucous neck cells
§  Secrete mucus
2.  Parietal cells
§  Secrete hydrochloric acid (HCl)
3.  Chief cells
§  Secrete the pepsin
4.  Enteroendocrine cells
§  Secrete hormones into the blood
§  G cells – Gastrin
§  Enterochromaffin (ECF) cells - histamine

Muscular, histologically diverse organ 30

 BIO270 L10
Intestines

§  Most nutrients are absorbed in the intestines

§  Four main layers
§  Mucosa
§  Villi and crypts
§  Submucosa
§  Blood and lymphatic vessels, nerves
§  Circular smooth muscle
§  Longitudinal smooth muscle

31
 BIO270 L10
Villi of intestine

§  Mucosal cells

§  Enterocytes
§  Absorptive cells with
microvilli
§  Goblet cells
§  Secrete mucus
§  Enteroendocrine cells
§  Secrete hormones
§  Paneth cells
§  Secrete antimicrobial
molecules
Also Stem cells – continually
replace epithelium
32
 BIO270 L10
Exocrine secretions from liver
Bile
§  Solution of digestive chemicals
and liver waste products
§  Produced in the liver and stored
in the gallbladder
§  Bile duct opens into small
intestine
§  Two main roles in digestion
1.  Digestion and absorption of fats
and fat-soluble vitamins
§  Emulsify fats
§  Amphipathic bile salts
2.  Elimination of hydrophobic
waste products
§  E.g. billirubin
33
 BIO270 L10
Exocrine secretions from the pancreas
§  Proteases break down protein
§  Secreted as inactive
proenzymes
§  Activated in the intestine
§  Amylase breaks down glycogen and
starch
§  Lipases break down triglycerides
§  Nucleases break down nucleic acids

§  Extracellular digestion in gut lumen

§  Digested products absorbed by
enterocytes

34
 BIO270 L10
BIO270 L10 Overview

Feeding and Digestion

1.  Overview
2.  Diet and energy
3.  Nutrient breakdown and transport
4.  Digestive systems
5.  Regulation of feeding and digestion

35
 BIO270 L10
Regulating feeding and digestion
§  Process of feeding requires energy and puts animal at risk
§  For many animals, feeding is regulated by the CNS
§  Signals sent to hypothalamus
§  Typically occurs when metabolic need can’t be met by circulating fuels

§  Digestive functions are controlled by nerve and hormone signals

§  Nerve signals
§  From central nervous system (CNS)
§  Within the gastrointestinal tract
§  Hormone signals
§  Wide variety of endocrine tissues and hormones
§  Three particularly important in humans:
§  leptin, peptide YY, ghrelin

36
 BIO270 L10
Control of Appetite

§  Three hormones control appetite by binding to

receptors in the hypothalamus
§  Leptin
§  Secreted by white adipose tissue when lipid content is high
§  Suppresses appetite
§  Peptide YY
§  Secreted by colon when full
§  Suppresses appetite
§  Ghrelin
§  Secreted by stomach when empty
§  Stimulates appetite

37
 BIO270 L10
Control of Appetite

§  Hypothalamic neurons release neurotransmitters in

response to hormones from the gut
§  Synapse with neurons from higher centers of the brain
to influence feeding behaviour:
§  Some neurotransmitters stimulate appetite
§  Neuropeptide Y (NPY)
§  Agouti-related peptide
§  Gamma amino butyric acid (GABA)
§  Some neurotransmitters inhibit appetite
§  Proopiomelanocortin (POMC)

38
 BIO270 L10
Hormonal control of Appetite

Control the
desire to eat
between meals

39
 BIO270 L10
Control of secretions

§  Hormones and neurotransmitters control digestive

secretions
§  acid
§  mucus
§  bile
§  bicarbonate
§  digestive enzymes

§  Respond to both the anticipation of food (perception

of prey) and its presence in the digestive system

40
 BIO270 L10
Control of gastric secretions

Parietal cell of stomach

HCl

41
 BIO270 L10
Control of Intestinal Secretions

Enters duodenum

Amino acids,
fatty acids

Vasoactive
Intestinal cholecystokinin
peptide

bicarbonate

Primary function of large intestine is reabsorption of water 42

 BIO270 L10
Control of gut motility
§  Food moves along the GI tract
by contractile waves of smooth
muscle
§  “Peristalsis”
§  Intrinsic myogenic activity
§  pacemaker cells (interstitial cells
of Cajal)
§  Receives signals from CNS
§  Motility controlled by myenteric
plexus
§  Optimal speed:
§  Fast enough to minimize
indigestible material in GI tract
§  Slow enough to allow time for
digestion and assimilation
§  Rate will vary according to diet
43
 BIO270 L10

Smooth visceral muscles along

digestive tract show rhythmic
cycles that move material along
tract through peristalsis

• peristalsis moves a bolus (or a

small oval mass of digestive
contents) along the length of the
digestive tract

44
 BIO270 L10
Metabolic transitions between meals
§  Postprandial period
§  Period immediately after feeding
§  Duration can vary from seconds to months
§  Nutrients are absorbed into the blood
§  Some nutrients are utilized and others are stored
§  Hormones control postprandial levels of nutrients
§  Insulin from β-cells of the pancreas
§  Stimulates glucose uptake and storage
§  Promotes conversion of glucose to fat
§  Glucagon from α-cells of the pancreas
§  Stimulates glucose release by liver
§  Glucocorticoids (e.g. cortisol) from adrenal cortex
§  Stimulates gluconeogenesis by the liver
§  Mobilize triglycerides from adipose tissue
45
 BIO270 L10
Metabolic Regulation in Insects

§  Hormones control

nutrient availability
§  Adipokinetic hormone
§  Secreted by the
corpora cardiaca
§  Stimulates release of
fatty acids from fat
body
§  Stimulates
breakdown of
glycogen into
trehalose

Figure 11.31
46
 BIO270 L10
Starvation response
§  Reorganization of metabolism to ensure long-term
survival
§  Goal: Conserve glucose to protect glucose-dependent
tissues; e.g. brain
§  Muscles shift to lipid metabolism

§  After lipid and glucose stores are depleted, protein

breakdown accelerates
§  Amino acids are converted to fatty acids and carbohydrates
§  Structural degradation of skeletal muscle occurs because there
is no protein store in the body

47
 BIO270 L10
Starvation response

48