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    Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

    PowerPoint Lectures forBio logy, Seventh Edit io n

    Neil Campbell and Jane Reece

    Lectures by Chris Romero

    Chapter 44

    Osmoregulation and

    Excretion

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    Overview: A balancing act

    The physiological systems of animals

    Operate in a fluid environment

    The relative concentrations of water andsolutes in this environment

    Must be maintained within fairly narrow limits

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    Freshwater animals

    Show adaptations that reduce water uptake andconserve solutes

    Desert and marine animals face desiccating

    environments

    With the potential to quickly deplete the body water

    Figure 44.1

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    Osmoregulation

    Regulates solute concentrations and balancesthe gain and loss of water

    Excretion

    Gets rid of metabolic wastes

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    Concept 44.1: Osmoregulation balances the

    uptake and loss of water and solutes Osmoregulation is based largely on controlled

    movement of solutes

    Between internal fluids and the external

    environment

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    Osmosis

    Cells require a balance

    Between osmotic gain and loss of water

    Water uptake and loss

    Are balanced by various mechanisms ofosmoregulation in different environments

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    Osmotic Challenges

    Osmoconformers, which are only marine

    animals Are isoosmotic with their surroundings and do

    not regulate their osmolarity

    Osmoregulators expend energy to controlwater uptake and loss

    In a hyperosmotic or hypoosmotic environment

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    Most animals are said to be stenohaline

    And cannot tolerate substantial changes inexternal osmolarity

    Euryhaline animals

    Can survive large fluctuations in external

    osmolarity

    Figure 44.2

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    Marine Animals Most marine invertebrates are osmoconformers

    Most marine vertebrates and someinvertebrates are osmoregulators

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    Marine bony fishes are hypoosmotic to sea water

    And lose water by osmosis and gain salt by bothdiffusion and from food they eat

    These fishes balance water loss

    By drinking seawater

    Figure 44.3a

    Gain of water and

    salt ions from food

    and by drinking

    seawater

    Osmotic water loss

    through gills and other parts

    of body surface

    Excretion of

    salt ions

    from gills

    Excretion of salt ions

    and small amounts

    of water in scanty

    urine from kidneys

    (a) Osmoregulation in a saltwater fish

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    Freshwater Animals Freshwater animals

    Constantly take in water from theirhypoosmotic environment

    Lose salts by diffusion

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    Freshwater animals maintain water balance

    By excreting large amounts of dilute urine

    Salts lost by diffusion

    Are replaced by foods and uptake across the gills

    Figure 44.3b

    Uptake of

    water and some

    ions in food

    Osmotic water gain

    through gills and other parts

    of body surface

    Uptake of

    salt ions

    by gills

    Excretion of

    large amounts of

    water in dilute

    urine from kidneys

    (b) Osmoregulation in a freshwater fish

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    Animals That L ive in Temporary Waters Some aquatic invertebrates living in temporary

    ponds Can lose almost all their body water and

    survive in a dormant state

    This adaptation is called anhydrobiosis

    Figure 44.4a, b (a) Hydrated tardigrade (b) Dehydrated tardigrade

    100 m

    100 m

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    Land Animals Land animals manage their water budgets

    By drinking and eating moist foods and byusing metabolic water

    Figure 44.5

    Water

    balance in

    a human

    (2,500 mL/day

    = 100%)

    Water

    balance in a

    kangaroo rat

    (2 mL/day

    = 100%)

    Ingested

    in food (0.2)

    Ingested

    in food (750)

    Ingested

    in liquid

    (1,500)

    Derived from

    metabolism (250)Derived from

    metabolism (1.8)

    Water

    gain

    Feces (0.9)

    Urine

    (0.45)

    Evaporation (1.46)

    Feces (100)

    Urine

    (1,500)

    Evaporation (900)

    Water

    loss

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    Desert animals

    Get major water savings from simple anatomicalfeatures

    Figure 44.6

    Control group

    (Unclipped fur)Experimental group

    (Clipped fur)

    4

    3

    2

    1

    0Wat

    erlostperday

    (L/100kgbodymass)

    Knut and Bodil Schmidt-Nielsen and their colleagues from Duke University observed that the

    fur of camels exposed to full sun in the Sahara Desert could reach temperatures of over 70C, while the

    animals skin remained more than 30C cooler. The Schmidt-Nielsens reasoned that insulation of the skin

    by fur may substantially reduce the need for evaporative cooling by sweating. To test this hypothesis, they

    compared the water loss rates of unclipped and clipped camels.

    EXPERIMENT

    RESULTS Removing the fur of a camel increased the rate

    of water loss through sweating by up to 50%.

    The fur of camels plays a critical role in

    their conserving water in the hot desert

    environments where they live.

    CONCLUSION

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    Transport Epithelia

    Transport epithelia

    Are specialized cells that regulate solutemovement

    Are essential components of osmotic

    regulation and metabolic waste disposal

    Are arranged into complex tubular networks

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    An example of transport epithelia is found in the

    salt glands of marine birds

    Which remove excess sodium chloride from theblood

    Figure 44.7a, b

    Nasal salt gland

    Nostril

    with salt

    secretions

    Lumen of

    secretory tubule

    NaCl

    Blood

    flowSecretory cell

    of transport

    epithelium

    Central

    duct

    Direction

    of salt

    movement

    Transportepithelium

    Secretory

    tubule

    Capillary

    Vein

    Artery

    (a)An albatrosss salt glands

    empty via a duct into the

    nostrils, and the salty solution

    either drips off the tip of the

    beak or is exhaled in a fine mist.

    (b) One of several thousand

    secretory tubules in a salt-

    excreting gland. Each tubule

    is lined by a transport

    epithelium surrounded by

    capillaries, and drains into

    a central duct.

    (c) The secretory cells actively

    transport salt from the

    blood into the tubules.

    Blood flows counter to theflow of salt secretion. By

    maintaining a concentration

    gradient of salt in the tubule

    (aqua), this countercurrent

    system enhances salt

    transfer from the blood to

    the lumen of the tubule.

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    Concept 44.2: An animals nitrogenous wastes

    reflect its phylogeny and habitat The type and quantity of an animals waste

    products

    May have a large impact on its water balance

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    Proteins Nucleic acids

    Amino acids Nitrogenous bases

    NH2

    Amino groups

    Most aquaticanimals, including

    most bony fishes

    Mammals, mostamphibians, sharks,

    some bony fishes

    Many reptiles

    (includingbirds), insects,

    land snails

    Ammonia Urea Uric acid

    NH3 NH2

    NH2O C

    C

    CN

    CO N

    H H

    C O

    NC

    HN

    OH

    Among the most important wastes

    Are the nitrogenous breakdown products ofproteins and nucleic acids

    Figure 44.8

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    Forms of Nitrogenous Wastes

    Different animals

    Excrete nitrogenous wastes in different forms

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    Ammonia Animals that excrete nitrogenous wastes as

    ammonia

    Need access to lots of water

    Release it across the whole body surface or

    through the gills

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    Urea The liver of mammals and most adult

    amphibians

    Converts ammonia to less toxic urea

    Urea is carried to the kidneys, concentrated

    And excreted with a minimal loss of water

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    Ur ic Acid Insects, land snails, and many reptiles,

    including birds

    Excrete uric acid as their major nitrogenous

    waste

    Uric acid is largely insoluble in water

    And can be secreted as a paste with little

    water loss

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    The Influence of Evolution and Environment onNitrogenous Wastes

    The kinds of nitrogenous wastes excreted Depend on an animals evolutionary history

    and habitat

    The amount of nitrogenous waste produced

    Is coupled to the animals energy budget

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    Concept 44.3: Diverse excretory systems are

    variations on a tubular theme

    Excretory systems

    Regulate solute movement between internal

    fluids and the external environment

    E P

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    Excretory Processes

    Most excretory systems

    Produce urine by refining a filtrate derived frombody fluids

    Figure 44.9

    Filtration.The excretory tubule collects a filtrate from the blood.

    Water and solutes are forced by blood pressure across the

    selectively permeable membranes of a cluster of capillaries and

    into the excretory tubule.

    Reabsorption.The transport epithelium reclaims valuable substances

    from the filtrate and returns them to the body fluids.

    Secretion.Other substances, such as toxins and excess ions, are

    extracted from body fluids and added to the contents of the excretorytubule.

    Excretion.The filtrate leaves the system and the body.

    Capillary

    Excretory

    tubule

    Filt

    rate

    Urine

    1

    2

    3

    4

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    Key functions of most excretory systems are

    Filtration, pressure-filtering of body fluidsproducing a filtrate

    Reabsorption, reclaiming valuable solutes from

    the filtrate

    Secretion, addition of toxins and other solutes

    from the body fluids to the filtrate

    Excretion, the filtrate leaves the system

    S f E t S t

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    Survey of Excretory Systems

    The systems that perform basic excretory

    functions

    Vary widely among animal groups

    Are generally built on a complex network of

    tubules

    P t h idi F l B lb S t

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    Nucleus

    of cap cell

    Cilia

    Interstitial fluid

    filters through

    membrane where

    cap cell and tubule

    cell interdigitate

    (interlock)

    Tubule cell

    Flame

    bulb

    Nephridiopore

    in body wall

    Tubule

    Protonephridia

    (tubules)

    Protonephr idia: F lame-Bulb Systems A protonephridium

    Is a network of dead-end tubules lackinginternal openings

    Figure 44.10

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    The tubules branch throughout the body

    And the smallest branches are capped by acellular unit called a flame bulb

    These tubules excrete a dilute fluid

    And function in osmoregulation

    M t h idi

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    Metanephr idia Each segment of an earthworm

    Has a pair of open-ended metanephridia

    Figure 44.11 Nephrostome Metanephridia

    Nephridio-

    pore

    Collecting

    tubule

    Bladder

    Capillary

    network

    Coelom

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    Metanephridia consist of tubules

    That collect coelomic fluid and produce diluteurine for excretion

    M l i hi T b l

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    Digestive tract

    Midgut

    (stomach)

    Malpighian

    tubules

    Rectum

    IntestineHindgut

    Salt, water, and

    nitrogenous

    wastes

    Feces and urineAnus

    Malpighian

    tubule

    Rectum

    Reabsorption of H2O,

    ions, and valuable

    organic moleculesHEMOLYMPH

    Malpighian Tubules In insects and other terrestrial arthropods,

    malpighian tubules

    Remove nitrogenous wastes from hemolymph

    and function in osmoregulation

    Figure 44.12

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    Insects produce a relatively dry waste matter

    An important adaptation to terrestrial life

    Vertebrate Kidneys

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    Vertebrate Kidneys Kidneys, the excretory organs of vertebrates

    Function in both excretion and osmoregulation

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    Concept 44.4: Nephrons and associated blood

    vessels are the functional unit of the

    mammalian kidney

    The mammalian excretory system centers on

    paired kidneys Which are also the principal site of water

    balance and salt regulation

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    Each kidney

    Is supplied with blood by a renal artery anddrained by a renal vein

    Figure 44.13a

    Posterior vena cava

    Renal artery and vein

    Aorta

    Ureter

    Urinary bladder

    Urethra

    (a) Excretory organs and major

    associated blood vessels

    Kidney

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    Urine exits each kidney

    Through a duct called the ureter

    Both ureters

    Drain into a common urinary bladder

    Structure and Function of the Nephron and Associated

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    (b) Kidney structure

    UreterSection of kidney from a rat

    Renal

    medulla

    Renal

    cortex

    Renal

    pelvis

    Figure 44.13b

    Structure and Function of the Nephron and AssociatedStructures

    The mammalian kidney has two distinct regions An outer renal cortex and an inner renal

    medulla

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    The nephron, the functional unit of the vertebrate

    kidney

    Consists of a single long tubule and a ball ofcapillaries called the glomerulus

    Figure 44.13c, d

    Juxta-

    medullary

    nephron

    Cortical

    nephron

    Collecting

    duct

    Torenal

    pelvis

    Renal

    cortex

    Renal

    medulla

    20 m

    Afferent

    arteriole

    from renalartery

    Glomerulus

    Bowmans capsule

    Proximal tubule

    Peritubular

    capillaries

    SEM

    Efferent

    arteriole from

    glomerulus

    Branch of

    renal vein

    Descending

    limb

    Ascending

    limb

    Loop

    of

    Henle

    Distal

    tubule

    Collecting

    duct

    (c) Nephron

    Vasa

    recta(d) Filtrate and

    blood flow

    F iltration of the Blood

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    F iltration of the Blood Filtration occurs as blood pressure

    Forces fluid from the blood in the glomerulusinto the lumen of Bowmans capsule

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    Filtration of small molecules is nonselective

    And the filtrate in Bowmans capsule is amixture that mirrors the concentration of

    various solutes in the blood plasma

    Pathway of the F iltrate

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    Pathway of the F iltrate From Bowmans capsule, the filtrate passes

    through three regions of the nephron

    The proximal tubule, the loop of Henle, and the

    distal tubule

    Fluid from several nephrons

    Flows into a collecting duct

    Blood Vessels Associated with the Nephrons

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    Blood Vessels Associated with the Nephrons Each nephron is supplied with blood by an afferent

    arteriole

    A branch of the renal artery that subdivides into the

    capillaries

    The capillaries converge as they leave the glomerulus

    Forming an efferent arteriole

    The vessels subdivide again

    Forming the peritubular capillaries, which surround theproximal and distal tubules

    From Blood Filtrate to Urine: A Closer Look

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    Proximal tubule

    Filtrate

    H2O

    Salts (NaCl and others)

    HCO3

    H+

    Urea

    Glucose; amino acids

    Some drugs

    Key

    Active transport

    Passive transport

    CORTEX

    OUTER

    MEDULLA

    INNER

    MEDULLA

    Descending limb

    of loop of

    Henle

    Thick segment

    of ascending

    limb

    Thin segmentof ascending

    limb

    Collecting

    duct

    NaCl

    NaCl

    NaCl

    Distal tubule

    NaCl Nutrients

    Urea

    H2O

    NaCl

    H2OH2OHCO3

    K+

    H+ NH3

    HCO3

    K+ H+

    H2O

    1 4

    32

    3 5

    From Blood Filtrate to Urine: A Closer Look

    Filtrate becomes urine

    As it flows through the mammalian nephronand collecting duct

    Figure 44.14

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    Secretion and reabsorption in the proximal

    tubule

    Substantially alter the volume and composition

    of filtrate

    Reabsorption of water continues

    As the filtrate moves into the descending limb

    of the loop of Henle

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    As filtrate travels through the ascending limb of theloop of Henle

    Salt diffuses out of the permeable tubule into theinterstitial fluid

    The distal tubule

    Plays a key role in regulating the K+and NaClconcentration of body fluids

    The collecting duct

    Carries the filtrate through the medulla to the renalpelvis and reabsorbs NaCl

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    Concept 44.5: The mammalian kidneys ability

    to conserve water is a key terrestrial adaptation

    The mammalian kidney

    Can produce urine much more concentrated

    than body fluids, thus conserving water

    Solute Gradients and Water Conservation

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    Solute Gradients and Water Conservation

    In a mammalian kidney, the cooperative action

    and precise arrangement of the loops of Henle

    and the collecting ducts

    Are largely responsible for the osmotic

    gradient that concentrates the urine

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    Two solutes, NaCl and urea, contribute to the

    osmolarity of the interstitial fluid

    Which causes the reabsorption of water in the

    kidney and concentrates the urine

    Figure 44.15

    H2O

    H2O

    H2O

    H2O

    H2O

    H2O

    H2O

    NaCl

    NaCl

    NaCl

    NaCl

    NaCl

    NaCl

    NaCl

    300

    300 100

    400

    600

    900

    1200

    700

    400

    200

    100

    Active

    transport

    Passive

    transport

    OUTERMEDULLA

    INNER

    MEDULLA

    CORTEX

    H2O

    Urea

    H2O

    Urea

    H2O

    Urea

    H2O

    H2O

    H2O

    H2O

    1200

    1200

    900

    600

    400

    300

    600

    400

    300

    Osmolarity of

    interstitial

    fluid(mosm/L)300

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    The countercurrent multiplier system involving

    the loop of Henle

    Maintains a high salt concentration in the

    interior of the kidney, which enables the kidney

    to form concentrated urine

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    The collecting duct, permeable to water but not

    salt

    Conducts the filtrate through the kidneys

    osmolarity gradient, and more water exits the

    filtrate by osmosis

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    Urea diffuses out of the collecting duct

    As it traverses the inner medulla

    Urea and NaCl

    Form the osmotic gradient that enables thekidney to produce urine that is hyperosmotic to

    the blood

    Regulation of Kidney Function

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    egu at o o d ey u ct o

    The osmolarity of the urine

    Is regulated by nervous and hormonal controlof water and salt reabsorption in the kidneys

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    Antidiuretic hormone (ADH)

    Increases water reabsorption in the distaltubules and collecting ducts of the kidney

    Figure 44.16a

    Osmoreceptors

    in hypothalamus

    Drinking reduces

    blood osmolarity

    to set point

    H2O reab-

    sorption helps

    prevent further

    osmolarity

    increase

    STIMULUS:

    The release of ADH istriggered when osmo-

    receptor cells in the

    hypothalamus detect an

    increase in the osmolarity

    of the blood

    Homeostasis:

    Blood osmolarity

    Hypothalamus

    ADH

    Pituitary

    gland

    Increased

    permeability

    Thirst

    Collecting duct

    Distal

    tubule

    (a)Antidiuretic hormone (ADH) enhances fluid retention by making

    the kidneys reclaim more water.

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    The renin-angiotensin-aldosterone system (RAAS)

    Is part of a complex feedback circuit that functionsin homeostasis

    Figure 44.16b

    Increased Na+

    and H2O reab-

    sorption in

    distal tubules

    Homeostasis:

    Blood pressure,

    volume

    STIMULUS:

    The juxtaglomerular

    apparatus (JGA) respondsto low blood volume or

    blood pressure (such as due

    to dehydration or loss of

    blood)

    Aldosterone

    Adrenal gland

    Angiotensin II

    Angiotensinogen

    Renin

    production

    Renin

    Arteriole

    constriction

    Distaltubule

    JGA

    (b) The renin-angiotensin-aldosterone system (RAAS) leads to an increase

    in blood volume and pressure.

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    Another hormone, atrial natriuretic factor (ANF)

    Opposes the RAAS

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    The South American vampire bat, which feeds

    on blood

    Has a unique excretory system in which itskidneys offload much of the water absorbed froma meal by excreting large amounts of dilute urine

    Figure 44.17

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    Concept 44.6: Diverse adaptations of the

    vertebrate kidney have evolved in different

    environments

    The form and function of nephrons in various

    vertebrate classes

    Are related primarily to the requirements for

    osmoregulation in the animals habitat

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    Exploring environmental adaptations of the

    vertebrate kidney

    Fi 44 18

    MAMMALS

    Bannertail Kangaroo rat

    (Dipodomys spectabi l is)

    Beaver (Castor canadensis)

    FRESHWATER FISHES AND AMPHIBIANS

    Rainbow trout

    (Oncorrhynchus mykiss)

    BIRDS AND OTHER REPTILES

    Roadrunner

    (Geococcyx cal i fornianus)

    Desert iguana

    (Dipsosaurus dorsal is)

    MARINE BONY FISHES

    Northern bluefin tuna (Thunnus thynnus)