EXERCISE 13  GASTROINTESTINAL MOTILITY AND SECRETION                          Afferent sensory nerve fibers
NEURAL CONTROL                                                                     Can be stimulated by
Enteric nervous system                                                                   o     Irritation of the gut mucosa
    2 plexuses                                                                          o     Excessive distention of the gut
          o     Myenteric plexus/Auerbachs plexus  outer, GI                           o     Presence of specific chemical substances in the gut
                movements, smooth muscle                                           Signals can cause excitation or inhibition of intestinal movements
          o     Submucosal plexus/Meissners plexus  inner, GI                     and/or intestinal secretion
                secretion and local blood flow, submucosa
    Can function on its own                                                   HORMONAL CONTROL
    Sensory neuron originate in GI epithelium or gut wall                     Hormones affecting GI motility
Myenteric plexus                                                                  Gastrin
    When stimulated                                                                     o   Secreted by G cells of the antrum of the stomach
          o     Increased tonic contraction of gut wall                                  o   Binds to receptors found predominatly on parietal and
          o     Increased intensity of contractions                                          enterochromaffin-like cells
          o     Slight increased rate of contractions                                    o   Released in response to stimuli associated with meal
          o     Increased velocity of impulse conduction, more rapid                         digestion
                movement of gut peristaltic waves                                                      Distension of stomach
    Not entirely excitatory                                                                           Products of proteins
    Send inhibitory signals to sphincter muscles                                                      Gastin releasing peptide  released by the
          o     Impedes movement of food along successive segment of                                    nerves of the gastic mucosa during vagal
                the GI                                                                                  stimulation
          o     Pyloric sphincter, ileocecal valve                                       o   Stimulated gastric acid secretion of parietal cells
Submucosal plexus                                                                        o   Stimulates growth of the gastric mucosa
    Function within inner wall of each minute segment of intestine               Cholecystokinin
    Sensory signals from GI epithelium integrate in submucosal plexus                   o   Secreted by I cells in the mucosa of the duodenum and
     to help control                                                                         jejenum
          o     Local intestinal secretion                                               o   Binds to receptors found on pancreatic acinar cells
          o     Local absorption                                                             (CCKA) and in brain and stomach (CCKB)
          o     Local contraction of the submucosal muscle (infolding of                 o   Released in response to digestive products of fat, fatty
                GI mucosa)                                                                   acids, and monoglycerides in the intestinal contents
Neurotransmitters secreted by enteric neurons                                            o   Stimulates secretion of pancreatic enzymes
    Acetylcholine  stimulates GI activity                                              o   Stimulates gallbladder contraction
          o     From parasympathetic postganglionic fibres                               o   Moderately inhibits stomach contractions  slows
    Norepinephrine and epinephrine  inhibits GI activity                                   emptying of food from the stomach to give adequate time
          o     From sympathetic pre- and post-ganglionic fibers                             for digestion of fats in the upper intestinal tract
    Others: ATP, serotonin, dopamine, cholecystokinin, substance P,              Secretin
     vasoactive intestinal polypeptide, somatostatin, leuenkephalin,                     o   First gastrointestinal hormone discovered
     metenkephalin, bombesin                                                             o   Secreted by the S cells in the mucosa of the duodenum
    Nerve endings                                                                       o   Released in response to acidification of the duodenum
Autonomic control of GI tract                                                            o   Mild effect on motility of GI tract
Parasympathetic innervation                                                              o   Stimulates pancreatic secretion of bicarbonate  in turn
                                                                                             helps neutralize the acid in the small intestine
    Cranial parasympathetic nerve fibers
          o     Almost entirely in the vagus nerves                               Gastrin inhibitory peptide
          o     Innervate esophagus, stomach, pancreas, first half of                    o   Aka glucose-dependent insulinotropic peptide
                large intestine                                                          o   Secreted by the mucosa of the upper small intestine
                                                                                         o   Released in response to fatty acids and amino acids but to
    Sacral parasympathetic nerve fibers
                                                                                             a lesser extent in response to carbohydrate
          o     Originate in spinal cord, pass through pelvic nerves
                                                                                         o   Decreases the motor activity of the stomach
          o     Innervate sigmoidal, rectal and anal regions
                                                                                                       Slows emptying of gastric contents into
          o     Function to execute defecation reflexes
                                                                                                        duodenum when the upper small intestine is
    Postganglionic neurons located mainly in myenteric and submucosal
                                                                                                        already overloaded with food products
     plexuses
                                                                                         o   Induce insulin secretion
    Stimulation of these nerves causes general increase in the activity of                            Stimulated by high glucose concentration in the
     the entire ENS  enhances activity of most GI functions                                            duodenum
Sympathetic innervation
                                                                                  Motilin
    Originate in the spinal cord between segments T-5 and L-2                           o   Secreted by the upper duodenum during fasting
    Innervate essentially all of the gastrointestinal tract                             o   Released cyclically and in response to alkaline pH in
    After leaving the spinal cord, most of the preganglionic fibers that                    duodenum
     innervate the gut pass the sympathetic chains and go out to outlying                o   Stimulates interdigestive myoelectric complexes
     ganglia                                                                                           Waves of gastrointestinal motility that move
    Most of the postganglionic sympathetic neurons originate from these                                through stomach and small intestine every 90
     ganglia, and travel to all parts of the gut                                                        minutes in a fasted person
          o     Preganglionic fibers  sympathetic chains  outlying                     o   Increases gastrointestinal motility
                ganglia  gut                                                            o   Controls the pattern of smooth muscle contractions in the
    Secrete norephinephrine and small amounts of epinephrine                                upper gastrointestinal tract
    Inhibits activity of the gastrointestinal tract                           Hormone           Secreted by         Stimuli for release     Major activities
    Effects exerted in 2 ways                                                 Gastrin           G cells of the    Meal ingestion          -stimulates gastric acid
                                                                                                 antrum of the       responses:              secretion of parietal cells
          o     Slight: direct effect of secreted norepinephrine to inhibit                      stomach             Distention of the      -stimulates growth of the
                intestinal tract smooth muscle (except mucosal muscle,                                                 stomach               gastric mucosa
                excited)                                                                                             Products of proteins
                                                                                                                     Gastric releasing
          o     Major: inhibitory effect of norepinephrine on the neurons of                                           peptide
                the entire enteric NS                                          Cholecystokinin   I cells in the    Digestive products of   -stimulates secretion of
    Stimulates the contraction of the GI sphincters                                             mucosa of the       fat, fatty acids, and   pancreatic enzymes
                                                                                                 duodenum and        monoglycerides in the   -stimulates gallbladder
    Stimulates vasoconstriction of the blood vessels associated w/                              jejenum             intestinal contents     constraction
     digestive system                                                                                                                        -inhibits stomach
    Strong stimulation can inhibit motor movements of gut  literally                                                                       contraction
                                                                               Secretin          S cells in the    Acidification of the    -stimulates pancreatic
     block movement of food through GI tract                                                     mucosa of the       duodenum                secretion of bicarbonates
                    duodenum                                   -mildly affects GI motility   EXERCISE 14  IN VITRO INTESTINAL ABSORPTION
Gastric             The mucosa of     Fatty acids and amino    -decreases the motor
inhibitory          the upper small   acids but to a lesser    activity of the stomach
                                                                                             Molecule        Transport into     Epithelial cells   Transport     Capillary
peptide             intestine         extent in response to    -induce insulin secretion                     epithelial cell    of villus          into
                                      carbohydrate                                                                                                 capillary
Motilin             Upper             Fasting state alkaline   -stimulates interdigestive
                    duodenum          pH in duodenum           myoelectric complexes
                                                                                                               Secondary
                                                               -controls the pattern of       Galactose          active
                                                               smooth muscle                   Glucose       transport with     Monosacchari        Facilitated
                                                               contractions in the upper                                                                          Enters
                                                               gastrointestinal tract
                                                                                                                   Na+                 des           diffusion
         Gastric secretion of stomach                                                                         Facilitated
                                                                                               Fructose
              o     Titrations  NSS washings, 2nd (2.2mL) and 3rd (2.9mL)                                      diffusion
              o     0.1M NaOH, assume 1.0mL HCl????                                                            Secondary
              o     pH = -log[H+]                                                               Amino            active
              o     Before and after stimulation of vagus nerve                                 acids        transport with
         Gastrointestinal motility                                                                                Na+
                                                                                                                                 Amino acids         Diffusion    Enters
              o     NSS vs atropine                                                                            Secondary
              o     Gavage with activated charcoal                                                               active
                                                                                               Peptides
              o     ACS                                                                                      transport with
              o     PDT                                                                                            H+
              o     Epinephrine                                                                 Short
                                                                                                                 Simple
                                                                                             chain fatty                        Short chain FA       Diffusion    Enters
                                                                                                                diffusion
HCl                                                                                             acids
         Produced in stomach by oxyntic/parietal cells                                                                          Triglycerides
                                                                                                                 Simple                                           Does
         Hydronium ions from rxn of water with carbon dioxide in parietal cells                Micelle                                 in           Lacteal
                                                                                                                diffusion                                        not enter
         Parietal cells release H+ ions together with Cl- ions                                                                  chylomicrons
         Vagal stimulation                                                                  Sugar absorption
               o     Decreased pH, more acidic                                                    Glucose  Na+ glucose symporter into epithelial cell, facilitated
               o     Vagal innervations in gastric glands containing the parietal                  diffusion (GluT2) out to capillary; Na/K pump
                     cells                                                                        Galactose  same as glucose
               1) Vagus nerve release Ach  stimulates gastric acid                               Fructose  facilitated diffusion through intestinal epithelium
                     secretion (neural)                                                           Pentoses  facilitated diffusion only, slow
               2) Enterochromaffin-like cells secrete histamine  stimulates                 Inhibitors
                     gastric acid secretion (paracrine)                                           DNP  inhibits ATP production, glucose transport not possible
               3) G cells stimulated by gastrin-releasing peptide  secrete                       Phlorhizin  displaces Na+ from binding site; glucose cannot bind to
                     gastrin  stimulates gastric acid secretion (endocrine)                       symporter, not transported
                                                                                                  Oubain/excess Li+ and K+ - inhibits Na+ pump
Gastric acid secretion                                                                       Intestine eversion
Increase                                     Decrease                                             Exposes highly-active mucosa to medium
Gastrin                                      Somatostatin (negative feedback)                     Stretching the small intestine increases the SA of the sac and
Cephalic and oral phases of a                Stress (sympathetic)                                  decreases sac wall
meal (parasympathetic)                                                                       Ringers solution with sodium bicarbonate
High levels of protein products              High levels of glucose and lipids                    Stimulates in vivo conditions and neutralizes pH drop, protects
Activation of parietal cells via Ach         Inhibition of Ach action                              intestine
Gastrin-relasing peptide                                                                     Aeration  for oxygen, molecule transport
Histamine                                                                                    Benedicts reagent
                                                                                                  Copper (II) blue + reducing sugar  copper (I) red
Histamine                                                                                         Effectiveness of glucose absorption
1) Activates adenylate cyclase                                                                    Higher glucose absorption, higher absorption
2) Increased intracellular cyclic AMP
3) Activation of protein kinase A                                                            Solutions
4) More efficient phosphorylation of cytoskeletal proteins (involved in                          Without DNP
     H+/K+ ATPase)                                                                                     o    glucose absorbed by intestine
5) More efficient release of H+                                                                        o    glucose left in the filtrate
                                                                                                       o    unreacted Benedicts
PDT = dist. charcoal*100length of intestine                                                           o    absorbance
   Greater PDT, greater distance travelled by substance                                               o   lower amount of glucose in solution
   Less inhibition of gastrointestinal motility by substance                                    With DNP
                                                                                                       o    glucose absorbed by the intestine
Atropine                                                                                               o    glucose left in the filtrate
    Competitive antagonist for muscarinic Ach receptors                                               o    unreacted Benedicts
    Blocks action of parasympathetic nervous system which decreases                                   o    absorbance
     intestinal motility                                                                               o   higher amount of glucose remained in solution 
    Should give low PDT value                                                                             absorption of glucose by the intestines was inhibited by
    PDT grp 1 (atropine) < grp 2 (NSS)                                                                    DNP
                                                                                                       o   due to uncoupling (proton leak from intermembrane space
Two tailed T-test                                                                                          to matrix)
   Atropine is similar to epinephrine
   Epinephrine                                                                                  Pentoses  simple diffusion
          o    Stimulates adrenergic receptors                                                   Glucose and galactose
          o    Causes relaxation, decrease in gut motility                                             o   Lumen to cell: SGLT-1, symporter with Na, active
          o    Stimulatory effect on sympathetic NS                                                        transporter dependent on Na-L ATPase pump in
          o    Inhibitory effect on parasympathetic NS                                                     basolateral membrane
                                                                                                       o   Cell to blood: GLUT-2, facilitated diffusion
                                                                                                 Fructose
                                                                                                       o   Lumen to cell: GLUT-5, facilitated diffusion
                                                                                                       o   Cell to blood: GLUT-2, facilitated diffusion
                                                                                                       o   Faster than pentoses but slower than other hexoses
EXERCISE 15  MEASURING CO2 PRODUCTION IN AQUATIC                                    Surface hypothesis  homeotherms maintain a constant temperature
ANIMALS                                                                               proportional to their body surface area
Direct methods of measurement                                                        Proportional rate of heat transfer between environment and body
     Calorimeter                                                                     SA,  loss of heat, will adapt =  metabolic intensity,  heat
     Hesss law                                                                      produced, RR
            o    Total amount of energy released in the breakdown of fuel            Larger animals have smaller surface areas
                 to a given set of end products is always the same                   SA of an object of isometric shape and uniform density varies as
            o    Heat released = chemical energy released                             0.67 (2/3) power of its uniform mass
Indirect methods                                                                     Mass increases as a cube of linear dimension, while SA increases
     Indirect calorimetry                                                            as a square
     Radioisotopes  atom movement tracking                                         When comparing different species, the exponent is closer to
     Respirometer  gas exchange                                                     approximately 0.75 (Kleibers law)
Basal metabolic rate
     Stable rate of energy metabolism measured in mammals and birds                 Weight-dependent  weight of the organism is directly involved or
      under the following conditions                                                  affects the metabolism
            o    Minimum environmental and physiological stress (at rest                   o     weight, metabolism
                 and with no temperature stress)                                     Surface-dependent  cell surface is presumably directly concerned
            o    Digestive and absorptive processes are temporarily                   with such energy requiring processes as the maintenance of ionic
                 deactivated (fasting)                                                disequilibria and the entry and exit of materials
     MR = aMb                                                                             o     SA, metabolism
     logMR = loga + b(logM)                                                         Intermediate  dependent on both weight and surface area
            o    where MR = basal metabolic rate
            o    M = body mass                                                   Errors
            o    A = intercept of the log-log regression line (differs between       High solubility of CO2, highly reactive
                 species)
                                                                                     Macromolecules can release CO2
            o    B = rate of change of MR with change in body mass
     Equivalent in ectotherms or poikilotherms are measured                     EXERCISE 16  OSMOREGULATION IN EARTHWORMS
            o    At controlled, specified body temp at which animal is not
                                                                                   Osmoregulation
                 expending additional metabolic energy to warm or cool                  o    Osmoregulators
                 itself                                                                 o    Osmoconformers
            o    Resting and fasting
                                                                                   Lumbricus
                                                                                        o    Euryhaline osmoregulator 2% NaCl, internal osmolarity
Body mass and metabolic rate
                                                                                             0.65% NaCl
   Small animals must respire at higher rates per unit body mass than                  o    Metanephridia and dorsal pores
    large animals
                                                                                        o    Collagenous cuticle layer  water cannot diffuse
   Inverse relation between the rate of O2 consumption per gram of
                                                                                   Metanephridia
    body mass and the total mass of the animal
                                                                                        o    Nephrostomes
   Inverse relation between metabolic rate and body mass applies                       o    Fluid isotonic to coelom, NaCl removed by active transport
    within species and between species                                                       system
Temperature and metabolic rate                                                          o    Majority of blood vessels with podocytes  coelom: major
   Poikilotherms  undergo temperature-induced changes in metabolic                         filtration
    rate                                                                                o    Enter metanephridia via nephrostome as coelomic fluid
   Homeotherms  expend metabolic energy to maintain a constant                        o    Narrow ciliated tubule: minor filtration process in blood
    body temperature                                                                         vessels
   Effect of temperature on reaction rate  temperature quotient is                    o    Brown ciliated & wide non-ciliated tubule: selective
    obtained                                                                                 reabsorption of water, protein and salts
   Temperature quotient  compares the rate of a reaction at two                       o    Bladder to nephridiopore: urine, urine excretion
    different temperatures                                                         Hyper-hyposmoregulation
   10 degrees Celsius is the standard temperature difference over                 Hyper-isosmoregulation
    which temperature sensitivity of a biological function is determined           Hyperosmotic environment
   Q10 = (K2/K1)^(10/(T2-T1))                                                          o    Hypersosmoreguation
                                                                                        o     reabsorption of water
    Pink  1mL 0.04% NaOH = 10mol CO2                                                                Concentrated urine
    RR = (mol CO2 A - mol CO2 B)/(weight of fish,grams x time,hrs)                   o     dorsal pore secretion
     temperature,  respiratory rate,  dissolved oxygen                                             moist surface
    When increased by 10 degrees Celsius, respiration doubles, Q10 =                   o    Aestivation
     2                                                                             Hypoosmotic environment
                                                                                        o    Isosmoregulation
Q10                                                                                     o     reabsorption fo water
   = 1, flat Q10, reaction rate is independent of temperature                                          concenctrated urine
   < 1, drops with increase in temperature                                                             volume
   > 1, increases with temperature                                                     o     dorsal pore secretion
                                                                                                        moist surface
Mass-specific metabolic rate                                                            o    out of burrows, soil surface
   Metabolic intensity
   Metabolic rate of a unit mass of tissue                                      EXERCISE 17  RENAL HANDLING OF WATER, SALT, AND
   O2 consumed per kg per hr                                                    SALICYLATE
   MR/M = AM(B-1)                                                                  Nephron = corpuscle + tubule
   (logMR)/M = logA + (B-1)logM                                                 A. Renal corpuscle  filtration
                                                                                         1. Bowmans capsule
Surface area to body mass relations                                                      2. Glomerulus
    Isometry  nonvarying proportions (same species)                                                a. Afferent arteriole
          o    Surface area varies as 0.67 power of biomass???                                       b. Efferent arteriole
    Allometry  systematic changes in body proportions with increasing          B. Renal tubule  reabsorption and secretion
     species size (different species)                                                    1. Proximal convoluted tubule
          o    Surface area varies as 0.63 power of biomass???                           2. Loop of Henle
                3. Distal convoluted tubule                                                                Directly increases Na reabsorption (PCT, loop of
                4. Collecting tubule                                                                        Henle, and principal cells of the late distal
           Vasopressin                                                                                     convoluted tubule)
                o    ADH                                                                                   Conserves fluid and salt (sodium); results in
                o    Pituitary gland                                                                        increase in blood volume and pressure
                o    Conditions                                                        Natriuretic peptides
                               Dehydration results in concentration of solutes              o    Natriuresis  excretion of sodium
                                plasma osmolarity increases                                   o    Heart and blood vessels
                               Large decrease in blood pressure (due to low                  o    Counteract effects of renin secretion, causing reduction of
                                water concentration)                                               blood pressure and extracellular fluid volume
                o    Functions                                                                o    BNP and ANP activated by atrial and ventricular
                               Increases water permeability of principal cells                    distension due to increase intracardiac pressure
                                and collecting ducts; increases urea                          o    Natriuretic and diuretic properties
                                permeability of inner medullary duct                                         Raise sodium and water excretion by increasing
                               Net effect  increases water absorption and                                   the glomerular filtration rate and inhibiting
                                conservation                                                                  sodium reabsorption by the kidney
                               Decrease in urine volume, higher concentration,        20% potassium chromate
                                lower osmolarity ?????                                 2.6% silver nitrate
           Aldosterone                                                                ferric chloride
                o    Mineralocorticoid produced in the adrenal cortex                  solving solving
                               Renin-angiotensin mechanism triggers release
                                of aldosterone
                               Juxtaglomerular apparatus
                               Renin ultimately catalyzes the production of           Fe + salicylate ion  complex, purple
                                angiotensin II, which stimulates aldosterone           Aspirin  salicylic acid (phenol) and acetic acid
                                release
                o    Function
Parameter          Extracellular Compartment                     Intracellular Compartment                 Total Body Water
                   Vol (L)     []          Total mOsm            Vol (L)     [ ] (mOsm/L) Total mOsm       Vol (L)      []                  Total mOsm
                               (mOsm/L)                                                                                 (mOsm/L)
Initial value      15          200         3000                  20         200          4000              35           200                 7000
Solution added     2           400         800                   0          0            0                 2            400                 800
Instantaneous                                                                                                           N.E.                7800
effect
After O.E.
Parameter          Extracellular Compartment                     Intracellular Compartment                 Total Body Water
                   Vol (L)     []          Total mOsm            Vol (L)     [ ] (mOsm/L) Total mOsm       Vol (L)      []                  Total mOsm
                               (mOsm/L)                                                                                 (mOsm/L)
Initial value      10          50          3000                  15                                        25
Solution added     0.5         100         800                   0          0            0
Instantaneous
effect
After O.E.