Feeding&

Food Processing

Feeding&

Food Processing1. Structure (review)

2. Function (behavior, physiology)

3. Nutritional needs

4. Digestive efficiency

1. Structure (review)

2. Function (behavior, physiology)

3. Nutritional needs

4. Digestive efficiency

Food captureFood captureFood captureFood capture

Mouth and pharyngeal cavity

JawsJaws

Teeth - jaw, mouth, Teeth - jaw, mouth, pharyngealpharyngeal

Gill rakersGill rakers

Mouth and pharyngeal cavity

JawsJaws

Teeth - jaw, mouth, Teeth - jaw, mouth, pharyngealpharyngeal

Gill rakersGill rakers

Fish Feeding - functionFish Feeding - function

Herbivores < 5% of all bony < 5% of all bony

fishes, no fishes, no cartilaginous fishescartilaginous fishes browsers - selective browsers - selective

- eat only the plant- eat only the plant grazers - less grazers - less

selective - include selective - include sedimentssediments

Detritivores 5 - 10% of all species5 - 10% of all species feed on decomposing feed on decomposing

organic matterorganic matter

Herbivores < 5% of all bony < 5% of all bony

fishes, no fishes, no cartilaginous fishescartilaginous fishes browsers - selective browsers - selective

- eat only the plant- eat only the plant grazers - less grazers - less

selective - include selective - include sedimentssediments

Detritivores 5 - 10% of all species5 - 10% of all species feed on decomposing feed on decomposing

organic matterorganic matter

Fish Feeding – functionFish Feeding – function

Carnivores zooplanktivoreszooplanktivores

suction feedingsuction feeding ram feedingram feeding

benthic benthic invertebrate invertebrate feedersfeeders graspersgraspers pickerspickers sorterssorters crusherscrushers

Carnivores zooplanktivoreszooplanktivores

suction feedingsuction feeding ram feedingram feeding

benthic benthic invertebrate invertebrate feedersfeeders graspersgraspers pickerspickers sorterssorters crusherscrushers

Fish Feeding – functionFish Feeding – function

More Carnivores fish feedersfish feeders

active pursuitactive pursuit stalkingstalking ambushingambushing luringluring

More Carnivores fish feedersfish feeders

active pursuitactive pursuit stalkingstalking ambushingambushing luringluring

Fish feeding behaviorFish feeding behavior

Fish feeding behavior integrates morphology with perception to obtain food:

Search --> Detection --> Pursuit Search --> Detection --> Pursuit --> Capture --> Ingestion--> Capture --> Ingestion

Fish feeding behavior integrates morphology with perception to obtain food:

Search --> Detection --> Pursuit Search --> Detection --> Pursuit --> Capture --> Ingestion--> Capture --> Ingestion

Feeding behaviorFeeding behavior

Fish show versatility in prey choice and ingestion

Behavior tightly linked to morphology

(co-(co-evolution)evolution)

Fish show versatility in prey choice and ingestion

Behavior tightly linked to morphology

(co-(co-evolution)evolution)

Fish feeding behaviorFish feeding behavior

Behavior tends to be optimizing when choices are available OptimalOptimal = maximize = maximize benefit:cost benefit:cost

ratioratio More for less!More for less! Select the prey that yields the Select the prey that yields the

greatest energetic or nutrient greatest energetic or nutrient “return” on the energy invested in “return” on the energy invested in search, pursuit, capture, and search, pursuit, capture, and ingestioningestion

Behavior tends to be optimizing when choices are available OptimalOptimal = maximize = maximize benefit:cost benefit:cost

ratioratio More for less!More for less! Select the prey that yields the Select the prey that yields the

greatest energetic or nutrient greatest energetic or nutrient “return” on the energy invested in “return” on the energy invested in search, pursuit, capture, and search, pursuit, capture, and ingestioningestion

Fish digestive physiologyFish digestive physiology

After ingestion of food, gut is responsible for: DigestionDigestion - breaking down food into - breaking down food into

small, simple moleculessmall, simple molecules involves use of acids, enzymesinvolves use of acids, enzymes

AbsorptionAbsorption - taking molecules into blood - taking molecules into blood diffusion into mucosal cellsdiffusion into mucosal cells phagocytosis/pinocytosis by mucosal phagocytosis/pinocytosis by mucosal

cellscells active transport via carrier moleculesactive transport via carrier molecules

After ingestion of food, gut is responsible for: DigestionDigestion - breaking down food into - breaking down food into

small, simple moleculessmall, simple molecules involves use of acids, enzymesinvolves use of acids, enzymes

AbsorptionAbsorption - taking molecules into blood - taking molecules into blood diffusion into mucosal cellsdiffusion into mucosal cells phagocytosis/pinocytosis by mucosal phagocytosis/pinocytosis by mucosal

cellscells active transport via carrier moleculesactive transport via carrier molecules

Digestive ApparatiDigestive Apparatitroutcarnivore

catfish

omnivore

carp

omnivore

milkfish

planktivore

Fish DigestionFish Digestion Two major groups: w/stomach, w/out w/out stomach: cyprinids (carps) w/stomach: cold-water salmonids, warm-

water catfish, tilapia, eels, grouper note: all “pure” predators have a stomach

and teeth relative gut length (RGL): gut:body length high RGL = species consuming detritus, algae

(high proportion of indigestible matter)

Two major groups: w/stomach, w/out w/out stomach: cyprinids (carps) w/stomach: cold-water salmonids, warm-

water catfish, tilapia, eels, grouper note: all “pure” predators have a stomach

and teeth relative gut length (RGL): gut:body length high RGL = species consuming detritus, algae

(high proportion of indigestible matter)

Relative Gut LengthRelative Gut Length

Species Feeding RGL

Labeo horie Algae, detritus 15.5

Garra dembensis Algae, inverts 4.5

Barbus sharpei Plants 2.8-3.1

Chelethiopselongatus

Zooplankton 0.7

Chela bacaila Carnivorous 0.9

Species Feeding RGL

Labeo horie Algae, detritus 15.5

Garra dembensis Algae, inverts 4.5

Barbus sharpei Plants 2.8-3.1

Chelethiopselongatus

Zooplankton 0.7

Chela bacaila Carnivorous 0.9

Fish Digestive Morphology: Major Divisions

Fish Digestive Morphology: Major Divisions

Mouth Esophagus Pharynx Stomach Intestine Rectum Secretory glands (liver and pancreas) often difficult to distinguish

Mouth Esophagus Pharynx Stomach Intestine Rectum Secretory glands (liver and pancreas) often difficult to distinguish

Gastrointestinal TractGastrointestinal TractGastrointestinal TractGastrointestinal Tract Esophagus Stomach

large in large in carnivores, small carnivores, small in in herbivores/omnivoherbivores/omnivoresres

Pyloric caeca Intestine

short in short in carnivores, long in carnivores, long in herbivores-herbivores-omnivoresomnivores

Anus - separate from urogenital pore

Esophagus Stomach

large in large in carnivores, small carnivores, small in in herbivores/omnivoherbivores/omnivoresres

Pyloric caeca Intestine

short in short in carnivores, long in carnivores, long in herbivores-herbivores-omnivoresomnivores

Anus - separate from urogenital pore

GI Tract- Secretory GlandsGI Tract- Secretory GlandsGI Tract- Secretory GlandsGI Tract- Secretory Glands Liver

produces bile produces bile (lipolysis)(lipolysis)

stores glycogenstores glycogen stores lipidsstores lipids

Pancreas digestive enzymesdigestive enzymes

proteases - protein proteases - protein breakdownbreakdown

amylases - starch amylases - starch breakdownbreakdown

chitinases - chitin chitinases - chitin breakdownbreakdown

lipases - lipid breakdownlipases - lipid breakdown

Liver produces bile produces bile

(lipolysis)(lipolysis) stores glycogenstores glycogen stores lipidsstores lipids

Pancreas digestive enzymesdigestive enzymes

proteases - protein proteases - protein breakdownbreakdown

amylases - starch amylases - starch breakdownbreakdown

chitinases - chitin chitinases - chitin breakdownbreakdown

lipases - lipid breakdownlipases - lipid breakdown

Digestive Anatomy: Mouth/Esophagus

Digestive Anatomy: Mouth/Esophagus

Channel catfish: large mouth/esophagus, capture prey, slightly predaceous, mouth has no teeth, no gizzard/cardiac sphincter

Common carp: small mouth for bottom feeding, pharyngeal teeth, grinds food

Tilapia: combination of bottom feeder, predator, efficient plankton feeder, uses gill rakers, pharyngeal mucous

Channel catfish: large mouth/esophagus, capture prey, slightly predaceous, mouth has no teeth, no gizzard/cardiac sphincter

Common carp: small mouth for bottom feeding, pharyngeal teeth, grinds food

Tilapia: combination of bottom feeder, predator, efficient plankton feeder, uses gill rakers, pharyngeal mucous

Digestive Anatomy: Stomach

Digestive Anatomy: Stomach

Channel catfish: have true stomach that secretes HCl and pepsinogen (enzyme)

Common carp: no stomach; however, “bulb” at anterior end of digestive tract, bile and pancreatic secretions empty into intestine posterior to cardiac sphincter, no secretion of gastrin (low pH)

Tilapia: modified stomach, secretes HCl, well-defined pocket, pH varies w/digestal flow, has pyloric sphincter

Channel catfish: have true stomach that secretes HCl and pepsinogen (enzyme)

Common carp: no stomach; however, “bulb” at anterior end of digestive tract, bile and pancreatic secretions empty into intestine posterior to cardiac sphincter, no secretion of gastrin (low pH)

Tilapia: modified stomach, secretes HCl, well-defined pocket, pH varies w/digestal flow, has pyloric sphincter

Digestive Anatomy: Intestine

Digestive Anatomy: Intestine

Channel catfish: length less than whole body, no large/small version, slightly basic pH, digestive secretions, nutrient absorption, many folds for absorption

Common carp: digestive tract is 3x whole body length, similar in activity to that of channel catfish

Tilapia: tract is 6-8x that of body length, activities similar to that of other species

Channel catfish: length less than whole body, no large/small version, slightly basic pH, digestive secretions, nutrient absorption, many folds for absorption

Common carp: digestive tract is 3x whole body length, similar in activity to that of channel catfish

Tilapia: tract is 6-8x that of body length, activities similar to that of other species

Digestive Anatomy: Liver and Pancreas

Digestive Anatomy: Liver and Pancreas

Both organs produce digestive secretions Liver produces bile but is also the primary

organ for synthesis, detoxification and storage of many nutrients

Pancreas is primary source of digestive enzymes in most animals

It also produces zymogens (precursors to enzymes)

Both organs produce digestive secretions Liver produces bile but is also the primary

organ for synthesis, detoxification and storage of many nutrients

Pancreas is primary source of digestive enzymes in most animals

It also produces zymogens (precursors to enzymes)

Fish Digestive PhysiologyFish Digestive Physiology Digestion is accomplished in

StomachStomach low pH - HCl, other acids (2.0 for low pH - HCl, other acids (2.0 for

some tilapia!)some tilapia!) proteolytic enzymes (mostly pepsin)proteolytic enzymes (mostly pepsin)

Digestion is accomplished in StomachStomach

low pH - HCl, other acids (2.0 for low pH - HCl, other acids (2.0 for some tilapia!)some tilapia!)

proteolytic enzymes (mostly pepsin)proteolytic enzymes (mostly pepsin)

Digestive Processes: Stomach

Digestive Processes: Stomach

Catfish as an example - its digestive processes are similar to that of most monogastric animals

Food enters stomach, neural and hormonal processes stimulate digestive secretions

As stomach distends, parietal cells in lining secrete gastrin, assisting in digestion

Gastrin converts the zymogen pepsinogen to pepsin (a major proteolytic enzyme)

Some fish have cirulein instead of gastrin

Catfish as an example - its digestive processes are similar to that of most monogastric animals

Food enters stomach, neural and hormonal processes stimulate digestive secretions

As stomach distends, parietal cells in lining secrete gastrin, assisting in digestion

Gastrin converts the zymogen pepsinogen to pepsin (a major proteolytic enzyme)

Some fish have cirulein instead of gastrin

Digestive Processes: Stomach

Digestive Processes: Stomach

Flow of digesta out of stomach is controlled by the pyloric sphincter

Pepsin has pH optimum and lyses protein into small peptides for easier absorption

Minerals are solubilized; however, no lipid or COH is modified

Mixture of gastric juices, digesta, mucous is known as chyme

Flow of digesta out of stomach is controlled by the pyloric sphincter

Pepsin has pH optimum and lyses protein into small peptides for easier absorption

Minerals are solubilized; however, no lipid or COH is modified

Mixture of gastric juices, digesta, mucous is known as chyme

Fish Digestive PhysiologyFish Digestive Physiology Digestion is accomplished in

StomachStomach IntestineIntestine

alkaline pH (7.0 - 9.0)alkaline pH (7.0 - 9.0) proteolytic enzymes - from pancreas & proteolytic enzymes - from pancreas &

intestineintestine amylases (carbohydrate digestion) - amylases (carbohydrate digestion) -

from pancreas & intestinefrom pancreas & intestine lipases (lipid digestion) - from lipases (lipid digestion) - from

pancreas & liver (gall bladder, bile pancreas & liver (gall bladder, bile duct)duct)

Digestion is accomplished in StomachStomach IntestineIntestine

alkaline pH (7.0 - 9.0)alkaline pH (7.0 - 9.0) proteolytic enzymes - from pancreas & proteolytic enzymes - from pancreas &

intestineintestine amylases (carbohydrate digestion) - amylases (carbohydrate digestion) -

from pancreas & intestinefrom pancreas & intestine lipases (lipid digestion) - from lipases (lipid digestion) - from

pancreas & liver (gall bladder, bile pancreas & liver (gall bladder, bile duct)duct)

Digestive Processes: Intestine

Digestive Processes: Intestine

Chyme entering the small intestine stimulates secretions from the pancreas and gall bladder (bile)

Bile contains salts, cholesterol, phospholipids, pigments, etc.

Pancreatic secretions include bicarbonates which buffer acidity of the chyme

Zymogens for proteins, COH, lipids, chitin and nucleotides are secreted

e.g., enterokinase (trypsinogen --> trypsin) Others: chymotrypsin, carboxypeptidase,

aminopeptidase, chitinase

Chyme entering the small intestine stimulates secretions from the pancreas and gall bladder (bile)

Bile contains salts, cholesterol, phospholipids, pigments, etc.

Pancreatic secretions include bicarbonates which buffer acidity of the chyme

Zymogens for proteins, COH, lipids, chitin and nucleotides are secreted

e.g., enterokinase (trypsinogen --> trypsin) Others: chymotrypsin, carboxypeptidase,

aminopeptidase, chitinase

Digestive Processes: Intestine

Digestive Processes: Intestine

Digestion of carbohydrates is via amylase, which hydrolyzes starch

Others: nuclease, lipase Cellulase: interesting in that it is

not secreted by pancreas, but rather produced by gut bacteria

Note: intestinal mucosa also secretes digestive enzymes

Digestion of carbohydrates is via amylase, which hydrolyzes starch

Others: nuclease, lipase Cellulase: interesting in that it is

not secreted by pancreas, but rather produced by gut bacteria

Note: intestinal mucosa also secretes digestive enzymes

Fish digestive physiologyFish digestive physiology Absorption is accomplished in

IntestineIntestine diffusion into mucosal cellsdiffusion into mucosal cells phagocytosis/pinocytosis by mucosal phagocytosis/pinocytosis by mucosal

cellscells active transport via carrier active transport via carrier

moleculesmolecules

Absorption is accomplished in IntestineIntestine

diffusion into mucosal cellsdiffusion into mucosal cells phagocytosis/pinocytosis by mucosal phagocytosis/pinocytosis by mucosal

cellscells active transport via carrier active transport via carrier

moleculesmolecules

Digestive processes: Absorption

Digestive processes: Absorption

Most nutrient absorption occurs in the intestine

Cross-section of the intestinal luma shows that it is highly convoluted, increasing surface area

Absorption through membrane is either by passive diffusion (concentration gradient)

Or by active transport (requires ATP) Or via pinocytosis (particle engulfed) Nutrients absorbed by passive diffusion

include: electrolytes, monosaccharides, some vitamins, smaller amino acids

Most nutrient absorption occurs in the intestine

Cross-section of the intestinal luma shows that it is highly convoluted, increasing surface area

Absorption through membrane is either by passive diffusion (concentration gradient)

Or by active transport (requires ATP) Or via pinocytosis (particle engulfed) Nutrients absorbed by passive diffusion

include: electrolytes, monosaccharides, some vitamins, smaller amino acids

Digestive processes: absorption

Digestive processes: absorption

Proteins are absorbed primarily as amino acids, dipeptides or tripeptides

triglycerides are absorbed as micelles COH’s absorbed as monosaccharides

(e.g., glucose, except for crustaceans) calcium and phosphorus are usually

complexed together for absorption all nutrients, excluding some lipids, are

absorbed from the intestine via the hepatic portal vein to the liver

Proteins are absorbed primarily as amino acids, dipeptides or tripeptides

triglycerides are absorbed as micelles COH’s absorbed as monosaccharides

(e.g., glucose, except for crustaceans) calcium and phosphorus are usually

complexed together for absorption all nutrients, excluding some lipids, are

absorbed from the intestine via the hepatic portal vein to the liver

Summary of Digestive Enzymes

Summary of Digestive Enzymes

Site/ Type Fluid/ enzyme Function/ notes

Stomach HCl Reduces gut pH,pepsiongen

Gastric secretions Zymogen, pepsinogen, HClAmylaseLipaseEsteraseChitinase

ProteolysisCOH’sLipidsEstersChitin

Pancreas HCO3

ProteasesAmylaseLipaseChitinase

Neutralizes HClCleave peptide linkagesCOH’sLipidsChitin

Liver/ bile Bile salts, cholestrol I ncrease pH, emulsifylipids

I ntestine AminopeptidasesLecithinase

Split nucleosidesPhospholipids to glycerol+ f atty acids

Site/ Type Fluid/ enzyme Function/ notes

Stomach HCl Reduces gut pH,pepsiongen

Gastric secretions Zymogen, pepsinogen, HClAmylaseLipaseEsteraseChitinase

ProteolysisCOH’sLipidsEstersChitin

Pancreas HCO3

ProteasesAmylaseLipaseChitinase

Neutralizes HClCleave peptide linkagesCOH’sLipidsChitin

Liver/ bile Bile salts, cholestrol I ncrease pH, emulsifylipids

I ntestine AminopeptidasesLecithinase

Split nucleosidesPhospholipids to glycerol+ f atty acids

Fish Nutritional NeedsFish Nutritional Needs

High protein diet:

Carnivores - 40 - 55% protein neededCarnivores - 40 - 55% protein needed Omnivores - 28 - 35% protein neededOmnivores - 28 - 35% protein needed

Birds & mammals - 12 - 25% protein Birds & mammals - 12 - 25% protein neededneeded

10 essential amino acids (PVT. TIM 10 essential amino acids (PVT. TIM HALL)HALL)

High protein diet:

Carnivores - 40 - 55% protein neededCarnivores - 40 - 55% protein needed Omnivores - 28 - 35% protein neededOmnivores - 28 - 35% protein needed

Birds & mammals - 12 - 25% protein Birds & mammals - 12 - 25% protein neededneeded

10 essential amino acids (PVT. TIM 10 essential amino acids (PVT. TIM HALL)HALL)

Fish Nutritional NeedsFish Nutritional Needs

High protein diet

Why so high? Proteins needed for growth of new Proteins needed for growth of new

tissuetissue Proteins moderately energy-dense Proteins moderately energy-dense

(don’t need dense source - ectotherms, (don’t need dense source - ectotherms, low gravity)low gravity)

Few side-effects - ease of NHFew side-effects - ease of NH44++

excretionexcretion

High protein diet

Why so high? Proteins needed for growth of new Proteins needed for growth of new

tissuetissue Proteins moderately energy-dense Proteins moderately energy-dense

(don’t need dense source - ectotherms, (don’t need dense source - ectotherms, low gravity)low gravity)

Few side-effects - ease of NHFew side-effects - ease of NH44++

excretionexcretion

Nutritional efficiency in fishes

Nutritional efficiency in fishes

Fish more efficient than other vertebrates:

Conversion factor = kg feed Conversion factor = kg feed required to produce 1 kg growth in required to produce 1 kg growth in fish fleshfish flesh

Fishes: 1.7 - 5.0Fishes: 1.7 - 5.0 Birds & mammals: 5.0 - 15.0Birds & mammals: 5.0 - 15.0

Fish more efficient than other vertebrates:

Conversion factor = kg feed Conversion factor = kg feed required to produce 1 kg growth in required to produce 1 kg growth in fish fleshfish flesh

Fishes: 1.7 - 5.0Fishes: 1.7 - 5.0 Birds & mammals: 5.0 - 15.0Birds & mammals: 5.0 - 15.0

Nutritional efficiency in fishes

Nutritional efficiency in fishes

Fish more efficient than other vertebrates

Why? Ectothermy vs. endothermyEctothermy vs. endothermy Energy/matter required to Energy/matter required to

counterbalance gravitycounterbalance gravity Bias of a high-protein dietBias of a high-protein diet

Fish more efficient than other vertebrates

Why? Ectothermy vs. endothermyEctothermy vs. endothermy Energy/matter required to Energy/matter required to

counterbalance gravitycounterbalance gravity Bias of a high-protein dietBias of a high-protein diet

Nutritional efficiencyNutritional efficiency

Maintenance ration (MR) = the amount of food needed to remain alive, with no growth or reproduction (% body wt./day)

MR is temperature-dependent MR increases as temperature increasesMR increases as temperature increases

MR is size-dependent MR decreases as size increasesMR decreases as size increases

Maintenance ration (MR) = the amount of food needed to remain alive, with no growth or reproduction (% body wt./day)

MR is temperature-dependent MR increases as temperature increasesMR increases as temperature increases

MR is size-dependent MR decreases as size increasesMR decreases as size increases