Chordates Part II Flashcards
aquatic mammals minimum body size
much larger than terrestrial
set by thermoregulatory demands of aquatic environment
larger mass animal bones
allometric growth
larger bones to support the weight, larger diameter, more robust
SA:V changes
as size increase, SA:V decreases
1unit cube = 6:1
2unit cube = 12:8 (1.5)
smaller ratio = lower rate of heat loss
sphere SA:V
SA = 4πr^2
V = (4/3)πr^3
smaller SA:V than cube of equal volume
minimize ratio for given volume
SA:V changes with shape
slender objects higher SA:V
ectotherms- lower MR, small, long, slender
consequences of size and shape variation
allometric relationships
eggs per female increase with body weight
influences survivorship and reproduction
evo devo
evolution and development
gene duplication
single genes
segment of chromosome
whole chromosome
whole genome
pseudogene
DNA sequences similar to normal genes but non-functional; as defunct relatives of functional genes
sub-functionalization
pairs of genes that originate from duplication, or paralogs, take on separate functions; ancestral gene-2 functions, new gene- 1 function
duplication events result in
pseudogenation
sub-functionalization
neo-functionalization
neofunctionalization
one gene copy, or paralog, takes on a totally new function after a gene duplication event; adaptive mutation process; one of the gene copies must mutate to develop a new function
functional divergence
genes, after gene duplication, shift in function from an ancestral function
gene duplications =
bursts of diversification
gene duplication, vertebrate evolution
3 episode widespread gene(ome) duplication
origin of verts, gnathostomes, teleosts
HOX clusters
4 in vertebrates
7-8 in teleost
snake venom toxins
co-opted from pancreatic origin
expanded by gene duplication
evolved under positive selection- neo-functionalization
Coqui development
no tadpole stage
rearrangment of development program
tail resorbed before hatching
adult characters (limbs) develop directly
frog with no direct development
tail growth before limb growth- gas exchange surface
classic neo-Darwinian 3-stage view of origin of species
mutation- new variant
selection- altered frequency/fixation (‘new population’)
reproductive isolation- new species
altered 4-stage evolved view of origin of species
mutation– new gene
re-programming- new ontogeny/individual
selection– new population
reproductive isolation– new species
re-programming
developmental/embryonic/ontogenic reprogramming or repatterning
mechanisms of developmental reprogramming
changes in developmental programs at various stages of life heterotopy heterochrony heterometry heterotypy
heterotopy
∆ location of gene expression
heterochrony
∆ timing of ≥2 processes relative to each other
- onset, offset, rate of process
- must be allometric
heterometry
∆ amount of gene product
heterotypy
∆ kind of gene product
transformation grid
1 species = reference
reference points relocated in derived species to reconstruct transformed grid
heterochronic change
∆ rate of development to maturity
∆ time to maturity
∆ time of onset of development
alone or combined, same or different times
classic neotony
axolotl- retains larval features (gills, fins)
neotony
paedomorphosis, retention of ontogenetic features into adulthood
heterochronic process graphs
∆ timing of development
a- ancestral, d- descendant, k- rate of shape development, a- rate of onset of growth, ß- age when offset shape is attained
paedomorphosis- development is truncated
deceleration
hyomorphosis
postdisplacement
paedomorphosis, deceleration
neotony
(-k): smaller slope, lower shape change in same time of development
paedomorphosis, hypomorphosis
(negative offset, progenesis)
same slope, shorter time period = smaller change in shape
paedomorphosis, postdisplacement
(positive onset)
onset of growth is later, offset is same, smaller change in shape
peramorphosis- development is extended
acceleration
hypermorphosis
predisplacement
peramorphosis, acceleration
(+k), steeper slope, faster change in shape over same time period, larger change in shape overall
peramorphosis, hypermorphosis
(positive offset)
start time same, end time later, longer period of development = greater change in shape
peramorphosis, predisplacement
(negative onset)
start time is earlier, end time is same, longer period of development = greater change in shape
shorter development time to maturity
miniaturization
either ∆ time to maturity or ∆ time of onset
∆ time to maturity
progenesis
∆ rate of development to maturity
neoteny
facultative paedomorphosis
environmentally induced polymorphism, results in coexistence of mature, gilled, fully aquatic paedomorphic adults and transformed, terrestrial, metamorphic adults in same population
really phenotypic plasticity
peramorphosis
individuals of a species mature past adulthood and take on hitherto unseen traits. It is the reverse of paedomorphosis
paedotypy
‘paedomorphosis’ but within a population- sometimes the organisms exhibit the change sometimes they do not
paedomorphosis in relation to paedotypy
comparison between species
descendents exhibit the change, ancestors do not
local heterochrony
changes in specific parts of body (animals are mosaics of different characters)
local terms- paedotypic somatic develop., per atypic gonadal develop.
why exhibit paedomorphosis
often determined by environment
saves energy of metamorphosis
early maturity
early reproductive output
amniote heart development
earlier development in all amniotes, not originally for endothermy, may be due to nature of egg– yolk movement, gas exchange
Tarsier
largest eye:body mass of all mammals
smaller than diapsids at initial devel.- allometric heterochrony
diapsid
(“two arches”) amniote tetrapods that developed two holes (temporal fenestra) in each side of their skulls
bird heterochrony
birds are miniature dinosaurs- pedomorphosis?
front limb:back limb larger in birds
birds have longer front limb relative to back limb
positive allometry of front limb
skull shape consistent w/ juvenile dino.
bird relaxed selection of front limbs
allowed them to ‘experiment’ with limb length- feeding?
led to wing development- exaptation
bird skulls
suggest pedomorphosis
retain juvenile shape overall and in bill, unlike dinosaur, alligator
bird mosaicism
peramorphic- bill, front limbs
paedomorphic- skull, back limb
ratites
ostrich, paedomorphic wing, skull; peramorphic hind limb, more robust skull
mosaic
mosaic animals
can’t say an animal is paedomorphic, must be more specific
giant anteater
very long snout, peramorphosis, allometric growth
bovid, kudu
very elaborate horns with large skull size
peramorphosis, allometric growth
peramorphosis in certopsian dinosaurs
bigger animal = larger differentiation from juvenile form
Hawaiin honeycreepers
peramorphosis in none, one, or both bills
human paedomorphosis
paedomorphic apes?
retention of younger developmental stages of apes
differential heterochrony between sexes
sexual dimorphism
blue boxfish: adult female is paedo. compared w/ male in body shape and color pattern
male anglerfish
salamander heterochrony
ovoviparous, viviparous
feeding much earlier in viviparous form
vivipary
development of the embryo inside the body of the mother
live birth
oviparous
animals that lay eggs, with little or no other development within the mother
ovoviparous
develop within eggs that remain within the mother’s body up until they hatch or are about to hatch
developmental trajectory
gradual, slow ontogeny or steps may be condensed for quicker ontogeny into fewer steps, then if one step is skipped you see bigger changes
salamander foot
B. occidentalis toes stop growing early, growth curve levels off, toes never project far out of pad— webbed foot, suction cup
timing of migration of neural crest cells
alters features, skin color
salamander- white = delayed crest cell migration- no color developed
color derivatives of neural crest cells
iridophores (blue?)
xanthophores (yellow)
erythrophores (orange)
melanocyte (black)
organ system
set of organs interacting to carry out major body functions
organ
body structure that integrates different tissues and carries out a specific function
vertebrate support/locomotion organ systems
skeleto-muscular system
vertebrate metabolism organ systems
respiratory system
digestive system
excretory system
vertebrae transport organ system
circulatory system
vertebrate reproduction organ system
reproductive system
vertebrate integration organ system
neuro-endocrine system (nervous system, endocrine glands)
vertebrate support and interaction organ
skin
homeostasis
maintaining stability, negative feedback
homeostasis feedback
environment ∆– physiological ∆– ∆ detected by neural receptors– info. sent along sensory pathway– integrator cells receive info. – info. sent along motor pathway– compensatory changes made by effector(s)– conditions returned to desirable levels
temperature regulation feedback
∆ detected by skin, hypothalamus– info. sent along afferent (sensory) pathway– neutrons receive sensory info. (brain)– info. sent along efferent (motor) pathway– actions
overall feedback model
increase/decrease– receptor (sensor)– integrator– effector(s)
nervous system main organs
brain, spinal cord, peripheral nerves, sensory orans, coordinates homeostasis
nervous systems present
in all metazoans except sponges
endocrine system organs
pituitary, thyroid, adrenal, pancreas, hormone-secreting glands
muscular system organs
skeletal, cardiac, smooth muscle- thermoregulation
skeletal system organs
bones, tendons, ligaments, cartilage
integumentary system organs
skin, sweat glands, hair, nails; skin largest organ, multiple functions
circulatory system organs
heart, blood vessels, blood; interacts w/ everything
lymphatic system
lymph nodes, lymph ducts, spleen, thymus
respiratory system organs
lungs, diaphragm, trachea, airways
digestive system organs
pharynx, esophagus, stomach, intestines, liver, pancreas, rectum, anus
excretory system organs
kidneys, bladder, ureter, urethra
reproductive system organs
ovaries, oviducts, uterus, vagina, mammary glands, testes, sperm ducts, accessory glands, penis
vertebrate coelom cavities
most have 2; pericardial (surrounding heart), pleura-peritoneal
mammals also have 2 pleural cavities (lungs)
coelom organs
organs are connected to cavity to be held in place
some organs outside of cavity (kidneys)
useless parts
vestigial, ‘hold overs’, ancestry
some human vestigial parts
third eyelid, darwin’s point, wisdom teeth, erector pili, body hair, coccyx, neck rib, thirteenth rib, fifth toe, paranasal sinuses, vomeronasal organ, fellowmen reflex, extrinsic ear muscles, subclavius muscle, palmaris muscle, plantaris muscle, pyramidalis muscle, appendix, male nipples, male uterus
third eyelid
Nictitating membrane- protects eye and sweep out debris, snow blindness, in birds, fish, amphibians, reptiles, tiny fold in inner corner of human eye
Darwin’s point
small, folded point of skin at top of ear in modern humans, remnant of larger shape to focus distant sound
wisdom teeth
early humans chewed lots of plants- another row of molars useful, only ~5% of population has a healthy set of 3rd molars
erector pili
smooth muscle fibres allow animals (mammals) to puff up fur to insulate or intimidate
- humans- goosebumps
- dogs/cats- fur standing up
body hair
brows- keep sweat out of eyes
male facial hair- sexual selection
most human body hair has no function
coccyx
fused vertebrae all that is left of tail
tail lost before humans began walking upright
neck rib
set of cervical ribs, leftovers from age of reptiles?, appear in <1% of population, cause nerve/artery problems, also associated w/ childhood cancer?
thirteenth rib
8% of adults have 13, most of us have 12
left over from chimps, gorillas?
fifth toe
mainly for balance in humans, grasping clinging to branches in apes
paranasal sinuses
nasal sinuses of ancestors may have been lined w/ odour receptors– heightened smell, aid survival
now- troublesome mucus-lined cavities, moistens air we breathe, makes head lighter
vomeronasal organ (VNO)
tiny pit on each side of nasal septum filled w/ nonfunctioning chemoreceptors
maybe once a pheromone detecting ability?
flehmen reflex
exposes VNO just behind front teeth (like horses)
expose to air, where pheromones are expected to be present
extrinsic ear muscles
trio of muscles, made it possible for pre hominids to move ears independently of heads
we still have them– ppl can wiggle ears
subclavius muscle
under shoulder from 1st rib to collarbone, useful for walking on all four
people have 0-2
palmaris muscle
long, narrow, runs from elbow to wrist, missing in 11% of humans, may have been for hanging, climbing
used for reconstructive surgery
plantaris
often mistaken for a nerve
useful for primate grasping with feet
not present in 9% of humans
pyramidalis
tiny, triangular, pouch like muscle, attached to pubic bone- from pouched marsupials?
>20% of humans don’t have
appendix
narrow, muscular tube, attached to large intestine for digesting cellulose when humans ate more plant matter, produces some white blood cells
>300,000 Americans/yr get it removed
male nipples
lactiferous ducts from well before testosterone causes sex differentiation in fetus
men have mammary tissue that can be stimulated to produce milk
male uterus
remnant of undeveloped female reproductive organ
hangs off male prostate gland
integument skin
injury, microbial, predator protection regulation of water regulation of Tb social interactions excretion/elimination of waste respiratory gas exchange muscle attachment sensory wrapping- shape and support
integument water regulation
water can pass both ways but amount that can pass varies in different animals- amphibians drink through skin
integument Tb regulation
hair, feathers, blood supply in skin, coloring
integument social interactions
color, size of feathers, chemical attractants from glands
skin characteristics
heaviest organ in body
most functions
remarkable repair functions
interface w/ environment, serious damage = serious problems
integument made up of
dermis and epidermis
dermis
lower layer
thick, protective functions
consists of layers
dermis made up of
stratum spongiosum stratum compactum hypodermis exoskeleton dermal plates/scales bone dentin(e), enamel chromatophores
stratum spongiosum
most of blood vessels that feed other layers of skin
stratum compactum
more compact layer below spongiosum
hypodermis
covering of muscles, fat deposits, muscles that allow skin to move relative to rest of body
dermis characteristics
collagenous and elastic fibres, fibroblasts, bones, scales, nerve fibres, blood vessels, smooth muscle, mesodermal
exoskeleton
reptiles, turtles, crocodiles
enamel
hydroxyapatite
less fibrous, harder (than bone or dentin)
fossil agnathans
ostracoderms
elaborate bony armour
derivatives of primitive dermal bone
lamellar bone
spongy bone
dentin
enamel
denticle
dentin + enamel
placoid shark scale
lamellar bone + dentin + enamel
kinds of bone
dermal/membane bone
endochondral bone
dermal bone
formed in membranes
intramembranous ossification
exoskeleton, dematocranium
endochondral bone
formed in cartilage
endochondral ossification
endoskeleton
chromatophores
dermis produced color, stellate, cells and pigment granules within move around
melanophores
liphophores
iridophores
stellate
neurons with several dendrites radiating from the cell body giving them a star shape
melanophores
contain melanin (dark pigment)
excess melanin
melanistic = black
lack of melanin
albinistic - very conspicuous, low survival
liphophores
contain corotanoids
xanthophores (yellow), erythrophores (red)
fossil dermis findings
skin pigments in extinct animals, convergence of melanism
ToF-SIMS to detect melanin
time-of-flight secondary mass spectrometry
composition & spatial distribution of surface molecules, including comparisons w/ spectra of melanin
SEM to detect melanin
scanning electron microscopy
presence of ovoid bodies consistent w/ melanophores
EDX to detect melanin
energy-dispersive x-ray microanlysis
carbon associated w/ skin and not adjacent sediment
evidence of melanism in 3 extinct animals
3 marine reptiles, each lineage secondarily aquatic
Ichtyopterygia, Mosasauroidea (Squamata), Eosphargis (Testudines, turtle?)
melanin function
thermoregulations- especially in turtle?
crypsis- ichthyosaur lacks countershading (deep diving habit, background matching in low light)
iridophores
contain crystal plates made of guanine- reflect light, influence perceived color
cyanophore
blue pigment, very rare, only known in a few species of fish
color changing
position of chromatophore
∆ distribution of pigment granules w/i chromatophore
seasonal moult
shifts in relative position of chromatophores
chromatophores- ameoboid
ex. if yellow pigments move onto of black pigments
distribution of pigment granules within chromatophore
densely packed or dispersed- density of color
seasonal moult
of plumage (birds) or pelage (mammals) color in epidermal structures can be 'dropped'
chameleons color changing
interactions- agressive, courting
antipredator response
dominant individual use color as social signal
sexual dichromatism
sexual dimorphism
greater in breeding season than rest of year- spend energy to enhance breeding color
seasonal color change
camouflage, varies geographically, may be shown some places and not others, background matching, moult btw color change
seasonal color change examples
Arctic Hare (Lepus arcticus) Rock Ptarmigan (Lagopus muta)
ontogenetic color change
color change through life, younger animals generally more vulnerable
ontogenetic change examples
mule deer- baby spotted, camouflage when laying down
Racer- snake, adults plain blue/grey, blotches on young
structural color
physical properties of body colouring
especially dramatic in birds
feathers refract light in various ways- differences in angle we look at it
blue amphibians
rare, usually not due to pigment, light is scattered by iridophores
chromatophore layers
filtering layer (xanthophore), scattering layer (iridophore), absorbing layer (melanophore) short λ (blue-green) largely absorbed by filter. med λ (yellow-green) pass through filter.- scattered by scattering layer- back through filter long λ (red-orange) - pass through filtering and scattering, absorbed by absorbing layer
parts of epidermis
stratum corneum
stratum germinativum
derivatives
stratum corneum
outer layer, shed old cells in flakes or one piece
stratum germinativum
below corneum, source of new cells which move up to outer layer
epidermis derivatives
various function glands keratinized structures (nails, claws, hooves, scales/scutes, hair, feathers, horns, antlers, foot pads, beaks)
glands
are IN dermis
BUT epidermal in origin
hair
dips down into dermis BUT epidermal derivative
keratin/lipids
barriers to water loss and UV
amounts of keratin are variable among taxa
mucus glands
moisture, gas exchange, cooling
granular glands
produce defence toxins
epidermal glands
mucous, poison, scent, sweat, sebaeous, mammary, uropygial
sebaceous gland
base of hair, lubricant for skin
uropygial glands
base of tail in birds, preening feathers, produces oil
mammarly glands
nipple- many ducts
teat- single duct
fish scales
bony scales, dermal, permanent, not shed, only lost through injury, persist throughout life, new growth every year
reptile scale
horny scales, epidermal, shed, called scutes
claw, beak, horn structure
central bony core, covered by vascularized dermis, outer epithelial layer
hair
keratonous, not modified scales, novel, grow from bone throughout life, multiple kinds (fine coat, second coat (guard hairs) grow through to provide protection)
feathers
down- close to body, small feathers, insulation
body contour feathers- grow through
flight feathers- moulted periodically and replaced, occur in tracks along body
skin as a sensory organ
touch receptors, transmitting pain, temperature, itch, touch information to CNS
important interface between body and environment
skin receptors
nociceptors, pruriceptors, thermoreceptors, mechanoreceptor, hair/glabrous skin, lips/tongue/cheeks, mystical pads, tactile foraging
nociceptors
pain
pruriceptor
itch
hair/glabrous skin reception
glabbrous/nebrous skin- free of hair (palms, soles)
discriminative touch- clearly distinguish differences in objects, descriminate more clearly
lips/tongue/inner cheeks reception
localization and movement of food
mystacial pads, vibrissae
snout of animals, long whiskers
vibrotactility, navigation, spatial orientation in dark
extend sensitivity beyond skin surface
tactile foraging
snout of star nose mole
elaborate w/ tentacles extremely sensitive to touch, finds way around and food
fish sounds
> 700 known vocal species
fish sounds
simple vs complex
same frequency, varying frequency/amplitude (moans, growls, peals)
how do fish make sounds
stridulation
air passage
drumming
stridulation (fish sounds)
rubbing/scraping together fins, bones, teeth
air passage (fish sounds)
little understood, internal movement of air, escape of air through mouth, gills, anus (farts), FRT- frequently repetitive ticks
waveform
amplitude vs. times
spectrogram
frequency (kHz) vs. time (s)
types of FRTs
3 types- FRT1, FRT2, FRT4
~2-8kHz, ~50-60dB, differ in amplitude
drumming (fish sounds)
‘sonic’ muscles pushing/pulling on internal air/swim bladder
males have longer muscles than females
why/when fish are vocal
spawning, courtship, agression, territorial, distress, predator/prey behaviour
cod drumming muscles
larger in males
larger at spawning time
correlated with fertilization potential
haddock courtship behaviour
pulse repetition rate changes at each stage of courtship- increases in frequency
studying fish sounds
passive acoustics
technology
passive acoustics
simply listening to sounds w/ hydrophones
non-invasive, non-visual (light not needed), continuous remote monitoring, provides detailed behaviour info
technology (fish sounds)
AULS
ROVs
Autonomous glider
AULS
autonomous underwater listening stations
ROVs
remotely operated vehicle
autonomous glider
buoyancy-drive AUV
moves through water independently, no engine, moves via density changes
ecological uses of fish sounds
locate vocal fishes
determine when fish are vocal
study of underwater noise effects
examine fish interactions
locating vocal fishes
identify essential fish habitat (EFH)
locate spawning habitats
exploration of the seas
census of marine life
determining when fish are vocal- season and time of day
spawning behaviour
predator/prey interaction
foraging
territorial defense
studying underwater noise effects
identify noise sources and levels
quantify temporal/spatial patterns in noise
quantify noise impact on fish behaviour
cusk-eel
found in Cape Cod by low tech passive acoustic methods, call in chorus just after sunset, tracks time of sunset through summer,
Haddock
using AULS 1000m deep, first in situ recordings in NA, recorded daily vocal activities- more vocal late in day, spawn mostly at night-
freshwater drum in hudson river
widely distributed highly vocal family, invasive, may spawn within canals that drain into Hudson
how did FW drum make it to hudson river
track acoustic path, with emphasis on spawning locations
found drum sounds in lake champlain canal, expected to spread dramatically and may alter rivers ecosystem
NEPTUNE canada subsea instruments
Hydrophones, seismometer, piezometer, bottom pressure recorder, gravimeter
piezometer
measure liquid pressure
gravimeter
measure local gravitational field
penetrometers
moisture, strength, harness of substrate
VENUS
Strait of Georgia, Saanich Inlet
UVic data archive, shore station, instrument platforms, nodes, autonomous vehicles, surface monitoring by BC ferries, satellites, gliders, profiling system
noisy ocean
peak listening is 1-10kHz (low frequency), lots of anthropogenic noise
fish hearing
fish have 2 inner ears, no middle or external ear, inner ear similar to other verts., sensory hair cells responsible for converting sound to electrical signal
potential effects on hearing
high intensity (transient)- fatigue, damage or kill sensory hair cells low intensity (shipping)- may have behavioural and physiological consequences
fish sensory cells
can be replace or repaired, unlike mammals
pile driving noise
direct mortality in surfperches
startle and alarm responses when exposed to air gun- rockfish, tighter school, school collapse, become motionless
behavioural effects of noise
distribution
fitness- reduced growth, reprod.
predator-prey interaction- interference
communication- range reduction, info loss
shipping noise
most extensive source of noise in ocean, especially along major shipping channels
reproductive consequences
physiological stress, restricting mate finding, keeping fish from preferred spawn sites
masking communicative sounds
impact ability of fish to communicate acoustically or use acoustic ‘soundscape’ to learn about envrionment
masking predator-prey relationships
affect ability to find prey or detect presence of predators
skeletomuscular system
vertebrate characteristic
internal, jointed skeleton (bone or cartilage)
works with muscular system
skeletomuscular functions
support of body movement via joints enclosure/protection of vital organs storage of minerals assistance in lung ventilation (amniotes)
skeletomuscular body support
ligaments, tendons, muscles
skeletomuscular mineral storage
Cap, P, Mg in bones
skeltomuscular lung ventilation
muscles connected to ribs
important connective tissues
cartilage bone ligaments tendons muscle
cartilage
matrix collagen chondroblasts chondrocytes lacuna(e)
chondrocytes
only cells found in healthy cartilage; produce and maintain cartilaginous matrix
chondroblasts
make cartilage matrix
lacuna(e)
hole in which cells grow
cartilage characteristics
more flexible than bone
most skeletons start w/ cartilage
offer support, bone growth
no blood vessels
types of cartilage
Hyaline
Fibrocartilage
Elastic
Hyaline cartilage
‘temporary’ cartilage during growth; most articulations, ribs, nose, larynx; least elastic; low collagen
Fibrocartilage
intervertebral disks, other joints (meniscus in knee); load bearing; show absorption; joint stabilization; able to resist pressure w/ minimum friction; moderately elastic; moderate collagen
Elastic cartilage
pinna, epiglottis, other parts of visceral skeleton; vibrational properties help emit/receive sound; most elastic; most collagen
knee minisci
important for knew function- load bearing, shock absorption, joint stabilization, joint lubrication, proprioception
proprioception
ability to sense stimuli arising within the body regarding position, motion, and equilibrium
bone properties
support and locomotion organic components mineral components mineral reserves dynamic
bone support and locomotion
balance between stiffness (hardness) and toughness (strength)
bone organic components
ex. collagen
toughness and elasticity
resistance to tensile loads
bone mineral components
ex. hydroxyapatite
stiffness, resistance to compressive loads
bone mineral reserves
Ca, P, Mg
bones, dynamic
modeling and remodelling
reabsorption and deposition
bone parts
osteoblast, osteocyte, osteoclast lacunae, canaliculi compact, spongy marrow woven, lamellar periosteum
osteoblasts
cells with single nuclei that synthesize bone
osteocytes
star-shaped cell, is the most commonly found cell in mature bone
osteoclasts
type of bone cell that resorbs bone tissue. This function is critical in the maintenance and repair, and remodelling of bones
bone lacunae and canaliculi
small canals between cells, blood cells and transport materials
periosteum
protective sheath around bones that connects to blood vessels and other structures like tendons
osteon
fundamental functional unit of much compact bone; bundle of blood vessels and lacunae
two types of tissue that form bone
compact
spongy
compact bone
cortical; facilitates bone’s main functions: to support the whole body, protect organs, provide levers for movement, store/release calcium; forms the cortex (outer shell) of most bones
spongy bone
cancellous, trabecular bone; higher SA:mass; less dense; softer, weaker, more flexible; suitable for metabolic activity-exchanges Ca; typically found at ends of long bones- proximal to joints, within interior of vertebrae; highly vascular; frequently contains red bone marrow- hematopoiesis
marrow
flexible tissue in interior of bones, 2 types yellow: fat red: blood cells birth- all red adult- 1/2 red
hematopoiesis
production of blood cells
woven bone
no uniform structure; early development; eventually replaced by lamellar bone
lamellar bone
compact, spongy, vascular canals, osteons, ‘plywood’ structure
plywood structure
regular parallel alignment of collagen into sheets (lamellae), mechanically strong, much lower proportion of osteocytes to surrounding tissue
bone stiffness
trade-off with toughness
high T low S: collagen– wood– chitin– bone– tooth dentin– mollusk shell– tooth enamel– glass, concrete, rocks, pottery
strain
dimensionless, epsilon = ∆length/length
stress vs. strain plot
elastic region– yield point– plastic region- fracture point
elastic region (stress vs. strain)
increases with high slope
rubber band like
steeper slope = less elastic
plastic region (stress vs. strain)
much lower slope increasing
stays together but is deformed
fracture point (stress vs. strain)
material breaks
stress
sigma = F/A
tissue stiffness =
y / x (stress/strain)
>yield pt. - yield or failure
bone elasticity
- 007% of strain
- 003 normal strain
- 015 results in fracture
ossification involves
direct or indirect
heterotropic bones
membranous ossification
direct laying down of bone- dermal armour, dermatocranium, parts of visceral skeleton, clavicle, others
endochondral ossification
indirect, cartilage precursor- most of axial and appendicular skeleton
ossification
laying down new bone material by osteoblasts- bone tissue formation
Heterotopic bones
isolated bones formed outside skeleton proper
sesamoid bones
small bones associated w/ tendons, joints; Often form in response to strain; act like pulleys, prove smooth surface for tendons to slide over increasing muscular forces
long bone structure
epiphysis, metaphysis, diaphysis
epiphysis
rounded end of a long bone, at its joint with adjacent bone(s)
diaphysis
the long midsection of the long bone
curious heterotopic bones
baculum, baubellum
Os penis, os clitoridis
baculum
penis bone, penile bone or os penis; bone found in the penis of many placental mammals, absent in human, function unknown- lock and key? some have projections, trident
baubellum
os clitoridis – a bone in the clitoris
bird bones
light skeleton, not necessarily light bones, hollow bones- air filled, not marrow filled; very dense bones, especially cranial compared w/ other animals
pneumaticity
air spaces in bones
post cranial pneumaticity
only birds, dinosaurs, perhaps gas exchange system
bone density
proportional to bone stiffness and strength
dense bone
stiffer, stronger, heavier
bone density vs. shape graph
heavy-light density vs. less-more rigid shape
min. density and rigidity = low stiffness and strength
max density and rigidity = high stiff. and strength
isoclines of stiffness and strength
medullary bone
woven bone, female birds, formed seasonally, prior to and during egg-laying, Ca reservoir for building hard eggshell
3 kinds of eggshells
hard
flexible
soft
hard shell
self-contained, rigid, fossils
calcareous matter dominates; tortoise, bird, dino, croc, gecko
flexible shell
needs water, calcareous layer loose, some fossils; turtles
soft shell
needs water, organic matter dominates, no fossils, gecko, tuatara, lizard, snake
crocodilian egg laying
pre-ovulatory hpercalcemia (takes 40% of Ca to make eggshells), no medullary bone formed
medullary bone significance
underscores evolutionary link btw. bird and dino
similar reproductive bio
means of sex ID in dino
ligaments
hold bones together, provide support, connective tissue, typically collagen
patellar ligament
between patella and tibia
holds tibia and femur together
2 main skeleton classifications
endoskeleton, exoskeleton
OR cranial, postcranial
exoskeleton
within integument
keratinized exo. - epidermis
bony exo. - dermis
endoskeleton
deep, within body
bony endo.
cartilagenous endo.
notochord
cranial skeleton
splanchnocranium
chondrocranium (cartilage)
dermatocranium
postranial skeleton
axial skeleton
appendicular skeleton
axial skeleton
vertebral column
notochord
appendicular skeleton
limbs
girdle
endoskeleton cartilage bone
vertebrae, ribs, limb bones
endoskeleton membrane bone
centra (teleost), sesamoid
exoskeleton dermal bone
skull roof, dentary, clavicle, gastrula, fish scales, osteoderm
gastralia
dermal bones found in ventral body wall of crocodilian/Sphenodon, between sternum and pelvis, do not articulate with vertebrae, support for abdomen, attachment sites for abdominal muscles
sphenodon
tuatara
main components of the skeleton
dermal
endoskeleton: somatic (axial, appendicular), visceral
median fin
median fin
one of the unpaired (i.e. dorsal, anal, and caudal) fins, restricted to fish, stability, propulsion
nuchal ligament
supports head, keeps it upright
degree of exoskeleton
greatly varies in all taxa
origin of vertebrate head skeleton
deep homology and co-option (exaptation)
spread of tissue through head (neural crest), not evolution of new skeletal tissue
axial skeleton
braincase, vertebral column, ribs
braincase
endochondral part of skull
vertebral column
backbone, tail, articulating vertebrae
first vertebra
atlas- allows up and down motion of head
atlas articulates with
occipital condyle(s) on back of braincase
second vertebra in amniotes
axis- allows rotary motion of head
parts of vertebra
centrum, neural arch and spine, zygapophyses (pre and post), diapophyses
occipital condyles
1 or 2 in tetrapods
undersurface protuberances of the occipital bone, articulates w/ superior facets of the atlas vertebra
centrum
main body of vertebra
neural arch
above centrum, spinal cord runs through
zygapophyses
projections of the vertebra that fit with adjacent vertebra; articulation, lateral/up/down motion, resist portion
diapophyses
the part of the transverse process of a thoracic vertebra that articulates with its corresponding rib
vertebrate lateral motion
many vert., including tetrapods, use lateral motion for locomotion, mammals- minimally
fish vertebral column
less flexible, without zygapophysis
dimetrodon
elaborate extension of neural spines, probably supported sail, evidence of vascularized tissue- thermoregulation, and/or social signalling
regionalizations of vertebral column
Trunk- Presacral, Cervical, Dorsal, Sacral
Caudal
Dorsal vertebra
thoracic, lumbar
frog vertebral column
very short, don’t bend well, highly reduced
mammal cervical vertebrae
7, typically do not have ribs
mammal ribs
thoracic vertebrae
mammal caudal vertebrae
tail, coccyx
urostyle
long bone-fused vertebrae at base of vertebral column, frogs and toads
bird vertebral column
stiff, lots of fusion, clavicle + inter clavicle = wishbone
wishbone
furcula, fusion of two clavicle bones
snake vertebral column
many vertebrae, large range of motion
autotomy
self amputation
lizard autotomy
fracture planes in vertebrae separate w/ muscle movement; tail moves back and forth rapidly, builds up lactic acid
regenerated tail is different- cartilaginous
Ribs
protect organs, used in breathing (muscle attachment), modified in various groups (ex. turtle)
tetrapod ribs
homologous w/ fish dorsal ribs
attached to sternum ventrally
reduction/loss (ex. anuran)
extras (ex. snakes)
fish ribs
dorsal/ventral/both
cobra
cervical ribs ‘spread out’ to give the illusion of being larger
Draco, lizard
wings, ribs articulate w/ vertebrae to spread out skin and form wings
rib newt
pierces own body wall w/ ribs to spread toxin
importance of sternum
not in animals that move ribs
very in animals that don’t, especially birds (keel)
appendicular skeleton
limbs, girdles
tetrapod pelvic girdle
firmly attached to sacrum- hind limbs need firm attachment to provide thrust (not in fishes)
sacrum
large, triangular bone at base of spine and upper, back of pelvic cavity, inserted between hip bones, number of fused vertebrae
number of vertebrae in sacrum
dogs- 3
humans, horses - 5
tetrapod pectoral girdle
not attached to head, often not attached to vertebral column (except in brachiators, flyers)
fishes- firmly attached to head
brachiators
primate, firm attachment of pectoral girdle for swinging
humerus, radius, ulna
present in crossopterygian- tetrapods- amniotes
deep homology, homologies in limbs
homologies
Similar characteristics due to relatedness
pelvic girdle bones
ilium, pubis, ischium
clavicle
present in fish and tetrapods- lost in some groups
only dermal element in mammal pectoral girdle
variable presence in mammals
mammal clavicle presence
present- human, bats
reduced- cats
most carnivores- absent or rudimentary
manus
carpals + metacarpals + phalanges
pes
tarsals + metatarsals + phalanges
variations in tetrapod manus and pes
homologies
convergences
fins, reduction/loss of fins, legs, ‘flippers’, wings, loss of flight, body elongation, reduction/loss of digits/limbs
functions of digits in tetrapods
support
locomotion
digging
grasping- perching, climbing, food manipulation
grasping behaviour in tetrapods
well conserved, chiefly arboreal life, feeding
very well-developed in tree frogs (manual and pedal)
best developed in mammals (manual and pedal)
grasping behaviour birds
front limbs modified to wings- grasping behaviour with back limbs, diverse toe configuration
grasping in lizards
negotiating complex habitat, varying degrees of manual and pedal grasping
grasping in unrelated tree frogs
convergence
suckers for gripping, digits that can wrap around branches
most bipedal tetrapod
birds, longer history of bipedalism that we do
modular locomotor system
hindlimb adapted for bipedal locomotion
shift from ab. muscle– back limbs for locomotor
shift from tail counter weight in dino.– knee as centre of gravity for straight back
theropod dinosaurs leading up to birds
all bipedal
joints
were bones meet, where all normal muscular function happens
kinds of joints
immoveable
slightly moveable
freely moveable
immoveable joints
synarthrosis
bones meet at a suture, associated w/ connective tissue
ex. skull bones
slightly moveable
amphiarthrosis
usually cartilage and connective tissue btw. bones, quite variable, ex. pubic symphysis (moves for child birth), spinal column
freely moveable
diarthrosis; synovial joint
subtypes: hinge joints, ball and socket joints, etc.
spinal column moveability
joints btw vertebrae, vertebrae move against each other but movement is limited
types of synarthrosis
serrate joint, scarf joint (wedge shape), butt joint (flush), peg and socket, lap joint (edges overtop each other, rare)
hinge joint
finger, knee, elbow; one-way movement
ball and socket joint
hip, shoulder; rotary motion
synovial fluid
reduces friction between articular cartilage of synovial joints during movement (freely moveable joints)
skull parts
chondrocranium
splanchnocranium
dermatocranium
chondrocranium origins
neurocranium, braincase- somatic
and neural crest
splanchnocranium origin
visceral skeleton, facial skeleton- from branchial arches
and neural crest
dermatocranium origin
skull roof- dermal
and neural crest
first gill arch
gnathostome jaw
second gill arch
hyomandiubular
jaw suspensions
amphistyly
hyostyly
autostyly
streptostyly
amphistyly
mandibular arch supported in part by hyomandibular, primitive Chondrichthyes
hyostyly
mandibular arch supported primarily by hyomandibula- Chondrichthyes, Actinopterygia
Autostyly
mandibular arch not supported by hyomandibule- Dipnoi, Tetrapoda
Steptostyly
quadrate bone moveable- Aves, Squamates
fate of meckel’s cartilage, arch 1
Articular (teleost, amph., reptile)— Malleus (mammals)
fate of palatoquadrate, arch 1
quadrate (teleost, amph., reptile)—- Incus (mammals)
fate of hyomandibula, arch 2
hyomandibula (teleost)– stapes (amph., reptile., mammal)
upper jaw teeth
mammals- restricted to 2 bones
other vets., more bones can support teeth
types of teeth
incisors- ripping
canines- stabbing
molars- chewing
carnivore dentition
incisors in front, large sharp canines, pointy triangular premolars, couple of molars
herbivore dentition
few small incisors, canines, space with no teeth, few premolars and molars- flat
omnivore dentition
teeth in same order but not much differences in shape/size, all relatively flat, fit together tightly, no spaces
temporal fenestration in amniotes
openings in side of skull, defined relative to position of bones; anapsid, synapsid, parapsid, diapsid
anapsid
ancestral, stem, lacking opening, early reptiles, turtles
diapsid
2 temporal fenestrae behind orbit, one superior and one inferior; dinosaurs, crocodilians, birds, tuaturas, lizards, snakes
synapsid
1 temporal fenestra behind the eye, below the postorbital bone, like the lower fenestrae in diapsids; extinct reptiles, mammals
parapsid
(euryapsids) extinct, ichthyosaurs, plesiosaurs; 1 fenestra behind the eye, above the postorbital, similar to upper fenestra of diapsids
streptostyly
quadrate bone rotates, increases mobility of jaws, lizards, snakes; 2 joints for jaw- one can be locked while other moves- more fore/apt movement, can swing out, can aid tongue projection, more forceful bite, can change in-lever
temporal fenestration in reptiles
sphenodon, crocs- unmodified diapsid
lizards- lower temporal bar lost- freeing quadrate
snakes- lower and upper bar lost, very open skull, highly developed streptostyly, more moveable quadrate
mammal temporal fenestration
temporal opening expanded and became confluent w/ orbital opening; bar btw eye and temporal fenestra lost
large open side of skull- large muscle attachment
turtle fenestration
anapsid but debatable- may be diapsid and secondarily anapsid
what is the function of temporal fenestrae
lighten skull without weakening, provide margins for muscle attachment, space for muscles to bulge out
pattern of temporal bar evolution in diapsids, especially squamates
lower temporal bar is lost very early in history of diapsids, is re-aquired in tuatara and others to reduce stress
function of derived lower bar in tuatara
reduction of stress on skull
squamate
scaled reptiles, are the largest recent order of reptiles, comprising all lizards and snakes
change from anapsid– diapsid
muscles from neurocranium to lower jaw (anapsid)– fenestra opens in dermatocranium– attachment of jaw muscles expands to edges of openings (therapsid)– jaw muscles attach to surface of dermatocranium (diapsid, synapsid)
zygomatic arch
cheek bone, zygomatic process of temporal bone- a bone extending forward from the side of the skull, over the opening of the ear
loss of lower temporal bar
allowed more musculature jaw– increases stresses on skull when animal bites– opened possibility for streptostyly
fixed quadrate
more stress on skull with biting
cranial kinesis
metakinesis, mesokinesis, prokinesis
movement of skull roof relative to braincase
metakinesis
joint between brain case and back of skull is at back of skull
mesokinesis
joint is in middle of skull- near orbits
prokinesis
joint in front of the orbit where snout articulates
symphysis
fibrocartilaginous fusion between two bones
evolution of snake gate
multiple joints all over skull, extremely mobile symphysis, not fixed like in humans, stretches
sphenodon skull
lower temporal bar, smaller jaw muscles and lower bite force than similar sized lizards, propalineal feeding, mastication (chew food more than other reptiles), handles food longer
croc. temporal bar
have bar, have strongest absolute bite of any living tetrapod; lizards the size of an alligator would have a much stronger bite (temporal bar lowers bite force)
propalineal feeding
close mouth- lower jaw in slightly posterior position– jaw slides forward- slides back in forth with food between teeth- temporal bar stabilizes jaw
secondary palate
found in various amniotes
best known in mammals
palate evolution
primary palate (early tetrapod)– growth across primary palate, shelf of bone (therapsid)– passageway btw primary and secondary palate, moves internal naris farther back into mouth, separate passage for eating and breathing (mammal)
internal naris
choana- the paired openings between the nasal cavity and the nasopharynx
succling mammal secondary palate
soft palate pressed against epiglottis- 2nd seal, allows swallowing milk and breathing (through nose)- disappears in adults because trachea drops
uvula
projection from posterior edge of middle of soft palate; almost completely unique to humans, unknown function and origin, involved in speech?
secondary palate bones
mammals- maxilla, premaxilla, palatine
crocs- those 3 + 1 more.. pterygoid?
why crocs have more elaborate secondary palate
flap closes off passageway for air, from water in mouth, so it can sit in the water for long periods of time ready to snap jaws shut
palate and stiffness
skull less resistant to bending if palate removed
maximum resistant with full palate
functions of muscular system
movement of body and parts, support, posture, protection of joints, internal transport, homeostatic adjustments, protein storage, metabolic heat production
muscular system internal transport
aids movements in blood vessels, digestive tract, reproductive tract
muscular system homeostatic adjustments
eyes- pupils constricting/dialating
muscular system heat production
shivering
blood flow musculature during before-during exercise
drastically change supply of blood to different body parts kidneys: 24% - 1% brain: 13% - 3% skin: 9% - 2% heart: 4.3% - 4.% skeletal muscle: 21% - 88%
types of muscle tissue
smooth, skeletal, cardiac
smooth muscle tissue
not striated, spindle shaped, not branched, involuntary, capable of slow sustained contractions, ex. walls of blood vessels
skeletal muscle tissue
striated, cylindrical, not branched, largely voluntary
cardiac muscle tissue
striated, cylindrical, branched, involuntary, looks like skeletal, ex. heart, involuntary- don’t control rate of heart beat, working all the time, branching propagates contractions
skeletal muscles
actin, myosin proteins- sarcomeres- make up muscle fibrils- make up muscle fibres– make up strap muscle
contraction of skeletal muscle
sliding of actin chains on myosin chains
shortened sarcomere length = increased overlap btw myosin and actin = maximum contraction = resting length
maximum force of a muscle depends on
being close to resting length
more x-sectional area- more potential force
velocity- max force at lowest velocity
length-tension curve of sarcomere
force vs. sarcomere length small sarcomere (hypercontracted)-- increasing up to max. force at resting length--- decreases to maximally extended sarcomere
speed of muscle contraction
- muscle configuration
- proportion of red and white fibre
- longer muscle can shorten more than shorter muscle
absolute muscle contraction
long muscle- more sarcomeres in series- can shorten more than a fibre with fewer sarcomeres in series
configuration of muscle fibres
parallel- strap, fusiform
pennate- angled (diff. angle than long axis)
bipennate- 2 different directions
pennate fibres
typically smaller, can fit into smaller places
muscle fibre cross section
anatomical, physiological
some dissipation of force if fibres aren’t in long axis direction, still contribute a lot of force
anatomical cross-section
across long axis of muscle
area of a slice through the widest part of the muscle perpendicular to muscles length
similar in parallel and pennate muscle
physiological cross-section
different in pennate b/c fibres are not parallel to long axis
area of a slice that cuts across all fibres of the muscle
different for a parallel and pennate muscle
tendons
connect muscle to bone, collagenous, all over the place, fairly elastic, can extend length by ~16%, store elastic energy when stretched which can be used by recoil to move body forward
muscle opperation
by contraction not relaxation
2 opposite actions need to take place (antagonism)
foramen magnum
hole at back of skull where spinal cord enters and connects w/ brain
acetabulum
hip, concavity, provides part of ball and socket joint w/ femur, head of femur fits into acetabulum
arm antagonism
extension: tricep contracts, bicep relaxes
flexion: tricep relaxes, bicep contracts
biceps and triceps are antagonistic
flight muscle antagonism
pectoralis- wing goes down
supracoracoideus- raises wing
synergism
perform ~same function in slightly different ways + up to ore complex action together
olecranon process
elbow, funny bone, where triceps connect
size depends on importance of tricep (ex. digging animal)
different size of antagonistic muscles
gull- downstroke more important
hummingbird- upstroke more important- larger supracoracoideus
categories of muscle function
extensor (extend), flexor (flex), adductor/abductor, levator, depressor, rotator, sphincter
adductor vs. abductor
adductor- bring body part towards body
abductor- takes body part away from body
levator vs. depressor
levator- raises
depressor- lowers
rotator
pronation- involves placing palms into the face-down position
supination- turns the palms anteriorly or superiorly to the supine (face-up) position
sphincter vs. dilator
sphincter- ringlike muscles surrounding and able to contract or close a bodily passage or opening
dilator- muscles that widen a body part
muscle insertion
typically stable end of muscle, sometimes more proximal part of muscle (closer to body)
muscle splitting and fusion
make homologies uncertain
axial musculature typically divided into
myomeres, separated by myosepta
myomere shapes
amphioxus: v-shaped
lamprey: w-shaped
shark- bony fish- more complexly folded
higher complexity - contraction extends beyond segment, important in locomotion
hypaxial and epaxial musculature
hypaxial- lie ventral to horizontal septum of vertebrae
epaxial- lie dorsal to the septum
amphibian/lizard motion
use lateral movement of body to extend stride
hard to move with limbs splayed to side
tetrapod motion
stride dependent on motion of limbs, musculature more developed around appendices, locomotory apparatus is limbs
snake axial musculature
expatiate use for contractions
remodelling muscle
hypertrophy, hyperplasia
hypertrophy
increasing size of individual muscle fibres
hyperplasia
increase in number of fibres, due to splitting of fibres
unused muscles
atrophy
snake, lizard reproductive modes
oviparity- lay eggs
viviparity- birth to live young with placenta
modes of delivery of nutrients to young
placentotrophy- delivery via placenta
lecithotrophy- delivery of nutrients via yolk- most reptiles (even viviparous)
process of forming yolk
vittelogenesis
income and capital of vittelogenesis
income- nutrients acquired to make yolk
capital- using previously stored nutrients
snakes more often use capital, female snakes often exhibit anorexia, don’t feed while carrying young- especially lose muscle (high protein store)
worlds smallest vertebrates
larval fish (<5mm)
larval fish characteristics
feed initially from yolk-sac
very poor swimmers
start with no vertebral column
stage of life history where recruitment is determined
larval fish mortality
> 99.9% - starvation, predation, advective losses (poor swimmers, carried away by currents in unfavourable conditions)
interannual variations in fish population abundance
<1915- variations in migration patterns
now know- due to recruitment
recruitment
variability in abundance results from interannual variability in # of individuals that survive larval stage
fisheries oceanography
branch of biological oceanography that studies the relationship between physical environment and abundance of marine fish
interannual variability in abundance reflects interannual variability in recruitment, proposed by
Johann Hjort, 1914
marine fish eggs
millions of eggs/ year
clear, buoyant, ~1mm diameter
preyed upon by zooplankton, larval fish, large fish (cannabalism common)
hatch times
days-months
colder water = longer time to hatch
yolk- sac
nutrition to developing embryo
aids in buoyancy
nutrients are function of mothers health
larval stage
2 phases: yolk-sac phase, post yolk-sac phase
large eyes, visual predators
suction feeding
yolk-sac phase
rely on yolk-sac, days-weeks (dependent on T), no gills no obvious fins, no proper tail
post yolk-sac phase
after yolk used up- exogenous feeding (plankton)
larva eat
initially copepod nauplii– switch to larger zooplankton
larval pray size depends on
foraging ability, gape (mouth width)
suction feeding
swim up to prey, open mouth quickly, creates vacuum, prey sucked into mouth
how far can larval fish see
about another body length away (~1cm)
prey are ~5cm apart, spend most of time foraging
reynolds number Re =
UL/v
U = swimming speed m/s
L = body length m
v = viscosity of seawater m^2/s - 10^-6 for 20º seawater
Re <100
viscous forces dominate, environment is totally viscous to animal, like human swimming through honey, larval fish in this range
Re > 200
intertial forces begin to dominate
typical reynolds numbers
sperm 0.01 copepod 4 larval fish 25 human 4x10^6 blue whale 3x10^8
reciprocal motion
fore-stroke and return are identical- useless in low Re conditions (must be non-reciprocating)
larval fish metamorphosis
transition from larval-juvenile
begins ~5-10mm
juveniles resemble miniature adults
mortality declines after metamorphosis
changes associated with metamorphosis
cutaneous (skin) breathing - gill breathing
develop paired pectoral fins, tail
develop adult-like pigment
eel-like swimming - beat and glide swimming
eye migration (flatfishes)
develop vertebrae- body rigidity for swimming
ELHS
early life history stages
ELHS atlantic cod
eggs 1mm
yolksac larva 3mm
late larval period 8mm
metamorphosed juvenile 10mm
c-start escape mechanism
larval fish, entire body curved like an eel or ‘C’ (no vertebral column)
as yolk-sac is absorbed and tail develops
swim speed increases response time decreases acceleration increases time to max speed decreases body curvature decreases
allometric growth in larval fish
head and tail grow relatively faster than rest of body- developing speed capabilities
after early development, change in growth rates
gills in fish
O2 uptake AFTER larval development (skin before)
ion exchange- Na+ uptake increases faster than O2 uptake
skin-gill transition
significantly earlier or Na+ uptake than O2 uptake
~16days vs. ~30?
ion exchange more important than respiration in larva?
why larval flatfish have eyes on both sides
living in 3D environment, need binocular vision (eggs are buoyant)
flatfish eye migration
can be as quick as 2 days, or 120
eyes kept in same plane as body turns
adaptation for 2D environment (ocean floor)
larval mortality graph
mortality %/day vs. length mm
- exponential
egg stage is highest percent and sharp slope
inflection point of graph is metamorphosis (~10mm)
metamorphosis distribution
tight distribution with size, not age- hydrodynamic constraints (Re number)
most undergo metamorphosis at 5-10mm
why metamorphosis is constrained by size
remodelling can’t be done at low Re
reciprocal motion doesn’t work at low Re
gill transition wouldn’t work at low Re
can’t have vertebral column in low Re (need flexibility)
fins no use in low Re (would move them back and forth)
fundamental units
mass, m, kg length, l, meter time, t, second force, F, newton work, W, joule power, P, watt
F =
m x a
W =
F x l
P =
W / t
types of muscular contraction
isotonic- concentric, eccentric
isometric
isotonic contractions
muscle changes length as it contracts- results in movement
concentric muscular contraction
force of muscle is adequate for moving a load
ex. picking up a stick
muscle shortens as it contracts
muscle contraction - sarcomeres
eccentric muscular contraction
muscle lengthens as it is contraction
ex. big heavy load you can’t pick up
isometric muscular contraction
muscle doesn’t change length as it contracts, constant length from one end to other including tendon connecting it, important in posture and support
ex. pushing a boulder you can’t move, pull open a door that won’t open
muscular force vs. speed
trade-off, decreasing, can’t maximize both at once, force is max at velocity = 0
force and power vs. shortening speed
force drops as velocity increases but power increases at intermediate velocity, can’t maximize force and power at the same time
classification of muscle fibres
fast-twitch fibres
slow-twitch fibres
some intermediates
fast-twitch fibres
white/blue, Type II; generate high force, rapid fatigue, high glycogen, anaerobic (glycolytic) metabolism- build up lactic acid, moderate blood and oxygen supply, low myoglobin, fast actions
slow-twitch fibres
red, Type I; low force, lower power, fatigue-resistant, abundant mitochondria- aerobic (oxidative) metabolism, myoglobin- transport hemoglobin, rich in blood and oxygen, can contract in sustained fashion
muscle fibre composition
speed depends on fibre composition, individual muscle can have both types of fibres, actions depend on amount of each type
duck breast muscle fibres
dark meat- red fibres- sustained flying
chicken breast muscle fibres
white meat- white fibres- can’t fly- fast twitch
force and power vs. velocity for slow and fast-twitch
slow-twitch- force has lower inflection point, power has lower max (~same as force infl. pt. in fast-twitch), max. velocity is ~1/2 that of fast-twitch
fast-twich have more power
contraction strength vs. time of muscles
eye: reaches max quick and dissipates quick- mostly fast-twitch
deep muscle of leg: reaches max slower and sustains it, declines much slower (mostly slow-twitch)
calf muscles: intermediate between the two
power performance and endurance
originally thought to not be a trade-off, after correcting for differences- found a negative correlation
can’t be a specialist and a generalist at the same time
endurance, sprint speed, lizards
high endurance = low sprint speed
high spring speed in ground dwelling- escape behaviour when entering open habitats, not seen in all lizards because its a trade-off
%red muscle in ocean species
large variations, constant swimmers = high proportion; benthic living = low proportion
position of red muscle, fish
usually superficial, internalized in tuna
power vs. tail-beat frequency, fish swimming
red muscle much lower in graph, much less powerful- slow, medium locomotion; white muscle kicks in and provides the power and fast locomotion
tuna vs. billfish
tuna: internalized red muscle, body remains stiff, caudal peduncle and tail are point of flexion
billfish: superficial red muscle, most of body involved in propagation of propulsion
senescence
muscular atrophy occurring with age, even if used; gradual deterioration of function
sarcopenia
degenerative loss of skeletal muscle mass (0.5–1% loss per year after the age of 50), quality, and strength associated with aging
whats going on with senescence
loss of fast-twitch fibres
shifting from fast-slow twitch phenotype with age, slowing of muscle contractile properties- reduces cost of locomotion in elderly
rattlesnake shaker muscles- rapid movement sustained for long periods of time
loaded with mitochondria and sarcoplasmic reticulum- supply Ca for nervous action
very economical- lowest cost per twitch
intermediate type of muscle
generates heat (one of the costs?)
levers
class 1, 2, 3 fulcrum between in-force and load in-force generated by muscular contraction lever-bone fulcrum- typically a joint speed/force depend on distances
class 1
out down, fulcrum, in up
ex. pushing down on toe, ankle, heel moving up
class 2
fulcrum, out up, in up
ex. pivot toe, leg goes up up, lift heal up
class 3
fulcrum, in up, out up
ex. pivot heel, push on leg, toe goes down
in-lever
l_i, length
out-lever
l_o
in-force
F_i
out-force
F_o, load
when in-force balances load
F_i * I_i = F_o * I_o
in-force moves load
F_i * I_i > F_o * I_o
load moves lever against in-force
F_i * I_i < F_o * I_o
steady state (levers)
F_o = F_i (I_i / I_o)
to increase F_o—- increase F_i, or I_i / I_o
digger vs. runner arm leaver
runner: short I_i, ratio is fairly low, not very big mechanical advantage; mechanical advantage tells a lot about function
to increase velocity of out-lever
decrease I_i / I_o
V_o * I_i = V_i * I_o
gear ratio
GR = I_o / I_i
low GR
power
high GR
speed (and stride in limbs)
direction of force of muscle
depends on orientation
arm at right angle- force directed along length of arm
arm open more than 90º- force ‘out’ from ‘elbow pit’
plantigrade
whole foot on ground- small metatarsals
digitigrade
walk on toes- med. metatarsals
unguligrade
walk on tips of toes- large metatarsals
orientation, speed of limb
effects gearing, speed
bear limb gears
high gear gluteal group- gluteus maximus, gluteus medius
low gear femoral group- adductor femoris
high AND low gear muscles used to extend femur
can rotate limb very rapidly with little power, rapid acting muscles, steady speeds
femoris- low gear for rapid acceleration
MA
mechanical advantage
Redeye Piranha
large adductor muscle, huge tendon, 3rd class lever, Li/Lo amplifies AM force transmission from jaw tip to posterior teeth- more powerful force at back of mouth
streptostyly in lizards, lever
2 different size in-levers and out-levers
upper articulation = longer in-lever = more forceful bite
cuticle
acellular outer mucus layer in fish, protective substance including toxins and antimicrobial compounds; limited keratinization
diversity of feeding types in fish
detritivores, planktivores, herbivores, carnivores, molluscivores, insectivores, piscivores, omnivores, parasites
evolution of feeding in fish
parasitize (jawless fish)– suction, biting (since jaw evolution, in most fish)
new mechanisms with bony fish
premaxilla protrusion, pharyngeal jaws, mechanical diversity, muscle duplication
important mouth functions
food capture- feet, mouth, teeth, tongue
oral transport- food handling in mouth, ingestion, mastication, swallowing, teeth, tongue, cranial kinesis, salivary glands
salivary glands
sublingual gland, mandibular gland, parotid gland, orbital gland; lubricate foods and start digestion
mobility of upper jaw
has evolved twice, led to increased processing capabilities, can tackle larger prey because they can break it into pieces as they kill it
grass carp pharyngeal jaws
no teeth in jaws, long serrated teeth in pharyngeal jaw- pharyngeal teeth; interact with basioccipital pad to grind down material making it more digestible
moray eel pharyngeal jaws
are brought forward when it opens its mouth and becomes an important prey capture mechanism- unable to generate pressure differences for suction feeding, massive adductor muscles propel pharyngeal teeth
feeding in water
prey is generally same density as water- approaching it pushes it away- most open mouth and oral cavity wide to create negative pressure- suck in prey and water
box turtle feeding
capable of feeding on land and in water (most turtle only in water), hyoid apparatus depresses more in feeding in water than in land
ways to swallow food whole
suction feeding, raptorial pharyngeal jaws, pterygoid walk, inertial feeding
suction feeding
teleosts, aquatic amphibians, aquatic turtles
raptorial pharyngeal jaws
moray eels
pterygoid walk
most snakes, move jaws independently over prey and pull it in
inertial feeding
birds, lizards, like a pelican
mechanical digestion
breaking food down into pieces
chemical digestion
in stomach
evolution of mammal chewing
- jaw joint, shapes of jaws changed so jaws be brought together to breakdown food unilaterally (one side of jaw at a time)
- change in jaw joint and adductor muscles- transverse movements (teeth can be moved side to side against each other)
- tribosphenic molars develop w/ complex surfaces, cusps that fit together dynamically during occlusion (can grind up food)
occlusion
manner in which the upper and lower teeth come together when the mouth is closed
tribosphenic chewing
unique to mammals, parallel to some dinos., puncture crushing- vertical bite first, then more side to side like horse/cow
increase lever arm of jaw muscles acting on jaw joint to increase chewing fores
moving muscle insertions further out on lower jaw
moving muscle insertions higher onto coronoid process
moving the position of the jaw joint to increase lever arm
arcilineal jaw movement
jaw closes, up and down, no fancy movement
propalineal movement
tuatara, jaws move against each other longitudinally
bird chewing
chew with guts not mouth, no teeth, beaks only for capture, can move both jaws, unique to raise upper jaw
gizzard
ventriculus- modified stomach, very muscular, horny sheet inside of it, keratinous sheet grinds up food
stomach stones in birds, to grind up food
gastroliths (typically rough rocks)
gizzard compensating for teeth loss
initially thought this, but these traits are seen together in some dinosaurs; probably aided reduction of head mass for flight
alimentary tract
tubular passage extending from the mouth to the anus, through which food is passed and digested
GI tract
gastrointestinal; esophagus, stomach, intestine; organ system responsible for consuming and digesting food, absorbing nutrients, expelling waste
sphincter GI tract
esophageal sphincter before stomach, gastric sphincter after stomach
gut regions
fore/mid/hind
cecum
beginning of large intestine; processing bacterial digestion of plant material, present in many verts.
parts of small intestine
duodenum, jejunum, ileum
parts of large intestine
cecum, colon, rectum, anus
changes in GI tract structure
straight- agnathan
spiral valve- chondrichthian
more and more complicated up to mammals
increasing surface area to improve digestion
the more plant material consumed
the longer the gut, difficulty with which plant material is digested
increasingly long and coiled intestines: carnivore- omnivore- hebivore
rumination
complex stomach with multiple chambers; regurgitate partially digested food from stomach (Cud), chew it again; Rumination- rechewing the cud, facilitates proper breakdown of cellulose rich plant matter
foregut
stomach, primary digestion, HCL
midgut
intestine, pancreas, liver; digestion, absorption, peptidases, amylases, etc.,
hindgut
hindgut chamber, rectum; absorption, defecation, fermentation
bird stomach(s)
proventriculus- secretes acids/enzymes
gizzard- mechanical breakdown
crop
dilation of esophagus that stores and softens food
gut lining
villi, which are lined with microvilli
enormously increase surface area
labile
to change
gut is labile
lots remodelling, increases in size with feeding, including increasing size of villi, increase seen in multiple organs (stomach, lungs, heart, pancreas, liver, kidneys, intestinal mucosa)
Hirschsprung’s Disease
Megacolon; musculature in gut stops working, faces are not moved properly, removed surgically
adaptive constipation
typical in large bodied vipers; may not deficit in 400days, provide balance when animal strikes, rapid strikes lunge it forward, retain feces more than other species that don’t lunge
atavism
resemblance to remote ancestors rather than to parents; reversion to an earlier type; ‘one-off’ developmental abnormalities, ‘throwbacks’; evolutionary reversals; problems for phylogenetic analysis
snake atavism
occasionally find a snake with Diddy biddy hind limb buds
human atavism
some babies born with tails
human coronary circulation similar to reptiles
Dollo’s law
biologist who argued that evolution can’t run backwards, genes/developmental pathways released from selective pressure will become nonfunctional
best example of evolution in reverse
axolotl- paedomorphosis lost, metamorphosis regained
viviparity in squamates, atavism
viviparity has evolved multiple times, most transitions are o-v, but in some cases v-o; if oviparity is ancestral (as is thought) then this represents a requisition
spontaneous atavisms
rare atavistic anomalies in individual specimens
phylogenetic character reversals
expressed in all members of a give clade
taxic atavisms
phylogenetic character reversals- important for evolution, mechanism for generating morphological variation within clades
atavisms and convergent evolution
can easily be confused if trees are equally parsimonious
double decay BS
double decay branch support
crocodilian atavism vs. convergence
similar long skinny snout- long thought to be convergent
molecular data shows sister species- snout derived- atavistic; skull table, braincase, jaws, hyoid, osteoderms, ribs, vertebrae, forelimbs, pelvis- reversals to fossil/outgroup traits
the case of the midwife toad
proteus with eyes restored
induced color adaptations by rearing on coloured soil
nuptial pads developed by forced water mating
nuptial pads
seasonal hypertrophy in skin of male frogs in water living species, hormonally controlled, help male keep grip on female for mating in water
hypertrophy
increase in the volume of an organ or tissue due to the enlargement of its component cells
hyperplasia
cells remain approximately the same size but increase in number
venom glands
modified salivary glands, venom kills prey, sometimes begins digestion
relative gizzard sizes
high fibre diet (hard to digest)- gizzard increases in size
low fibre- gizzard decreases in size
gizzard varies between and within species, gut readily remodelled
respiratory gas exchange
oxygen gain from fluid medium, CO2 dumped into fluid medium
ventilation
movement of medium (water/air) either due to current or muscular action on a part of the animal, especially in relation to the gas exchange surface
breathing
skeletoventrical movements that cause ventillation near the gas exchange surface
respiratory gas exchange organs
gills, lungs, skin
skin for gas exchange
majorly amphibians but to some degree in all animals, even a little bit in humans
plethodontid salamanders
loss of lungs
loss of larval stage
consequences in tongue projection
plethodont lung loss
synapomorphy, ancestral character, anti bouyancy mechanism, changes in breathing- lose need for hyoid apparatus (movements of mouth floor)
ancestral lung state
salamanders lived in fast flowing streams- high O2
not the case now, lung loss is not a function of O2
plethodont, loss of larval stage
direct development, in some species, no requirement of hyoid for suction feeding
plethodont tongue projection
ballistic tongue projection; hyoid apparatus projected out of mouth (tongue skeleton), retracted by muscles all the way back to hip; only possible b/c hyoid not needed for buccal pumping
O2 concentrations
fresh water 6.6 mL/L at 20ºC
Air 209 mL/L
increases with declining temperature and increasing turbulance
increasing skins respiratory exchange
loose, baggy skin, increased SA (hellbender salamander, lake Titicaca frog), capilli growth- highly vascularized gas exchange surface (male hairy frog)
larval salamander gas exchange organs
skin, lungs, gills
bony fish gas exchange organs
lungs very basal, secondarily lost in many groups, modified into swim bladders in many species- triple exaltation (breathing, buoyancy, sound?)
gills
main aquatic gas exchange surface, fish, amphibians
pharyngeal arch- gill arch- skeletal support for the gill
on each gill arch (gas exchange)
primary lamelli, covered in secondary lamelli- these are the actual gas exchange surface
counter current exchange system, gills
water flows across secondary lamella on gill arch, they pick up oxygen from water, and carry the oxygen to body tissues in the opposite direction of water flow
lamprey water flow
nonfeeding: through mouth– pharynx– gill arches– out
feeding: in sides of gill arches and back out, doesn’t enter body cavity, mouth, or pharynx
spiracle
opening in sharks where water enters and can be forces out gill slits
operculum
bony flap covering gills, can be closed
teleost fish respiratory (gills)
take in water in buccal cavity with operculum closed– expand opercular cavity, pressure drops (same as feeding)– force water through opercular cavity– opercular valve open– water out
salamander gill
remain external, well developed
frog spiracle
dictates direction of water flow through gills (tadpole), types 1,2,3,4
variation in gill sizes
large gills- still ponds
small gills- fast flowing mountain stream
bigger fin on tail- pond
larger gas exchange surfaces
lungs
major gas exchange surface in air
gills have too many fine surfaces, would not be efficient in air
evolution of aspiration breathing in tetrapods
- aquatic buccal pump- operated by hyoid apparatus
- two-stroke buccal pump- 2 movements of mouth for each breath, sole dependence on buccal pump
- exhalaion powered by hypaxial musculature
- costal aspiration (loss of buccal pump, fully associated with musculature)
aspiration
bringing in air via musculoskeletal system- sucking in air
2-stroke buccal pump
drops floor of mouth to open mouth cavity (using buccal pump)– then glottic opens– air is forced out past air that has just been taken in– floor of mouth raised– air forced past lungs
frog/amphibian lung breathing
breathe through nose– glottis closed– open nostril– lower floor of mouth– negative pressure– air enters oral cavity– open epiglottis– force air out nostril (exhale) by elastic recoiling of lung– close nostril– raise floor of mouth (second stroke)– force air into lung- set up elastic recoil
epiglottis
a thin, valvelike, cartilaginous structure that covers the glottis during swallowing, preventing the entrance of food and drink into the larynx
glottis
opening between the vocal cords at the upper part of the larynx
in between breathes, frogs
raise and lower floor- get rid of stale air in mouth
frog courtship noises
with nostrils closed- force air into vocal sacs rapidly- accoustic radiator- shift air back and forth between lungs and vocal sacs very rapidly
sprawling posture and breathing
body musculature needed for locomotion, breathing?
volume moved out of lungs decreases rapidly with speed, can’t run fast for long- can’t breathe- trade-off
minute ventilation
total air inhaled and exhaled in a minute
axial constraint
breathe and move with same musculature
gular pump
accessory breathing apparatus- independent of body musculature so they can move air into lungs while running
oropharyngeal pump used for lung inflation in air-breathing fishes and amphibians
buccal pump
pharyngeal pump used as accessory lung inflation mechanism in lizards and tuataras
gular pump
non-ventilatory expansion/compression of buccal cavity, preformed with mouth closed and usually serving as olfactory function
buccal oscillation
non-ventialtory expansion/compression of buccal cavity, with mouth open, serving as thermoregulatory function
gular flutter (related to panting)
Ichthyostega
had a rib cage, perhaps breathing close to amniotes
elastic recoil
exhalation
recoil aspiration
lung wall musculature contracts– pressure drops– lungs deflate, air pushed out– integument drawn inward to compensate for volume change— deformation stores elastic energy - negative pressure
tetrapod respiratory system
tidal or unidirectional
dead space
vocalizations
tidal respiration
bidirectional, humans
unidirectional
birds, more efficient O2 extraction
dead space
volume of air inhaled that does not take part in gas exchange, because it (1) remains in the conducting airways, (2) reaches alveoli not perfused
perfused
supply (an organ, tissue, or body) with a fluid, typically blood, by circulating it through blood vessels
benefits of dead space
CO2 retained, make buffered blood; Inspired air brought to T_b, increasing affinity of hemoglobin for O2, improving uptake; Particulate matter trapped on mucus, allowing removal; Inspired air is humidified, improving quality of airway mucus
vocalization
vocal chords- in larynx, vibrate when air rushes past
syrinx- vocal organ of birds; at the base of trachea, produces sounds w/o vocal cords, sound is produced by membrane vibrations when air flows through
types of lung
faveolar lung
alveolar lung
faveloar lung
septate, reptiles, modified in birds, less compartmentalized, no alveoli, pockets open from central chamber
alveolar lung
mammals, lots of alveoli pickets
compliance
ability for the lung to be inflated
parenchyma
gas exchange tissue
structural type vs. praenchyma
uni-cameral, multi-cameral, highly specialized vs. homogeneous, heterogeneous
highly specialized, homogeneous
large surface area, low compliance, mammal
highly specialized, heterogeneous
large surface area, high compliance, dinosaurs, birds
uni-cameral, homo-heterogeneous
amphibians, reptiles, large surface area, low compliance unless body elongated
amphibian lungs
single chambered, only complement gills and skin
amniote lungs
multichambered shared by all amniotes, principle gas exchange site, key to conquering land
Archosaurs
crocodiles, birds
Lepidosaurs
lizard, snakes, tuatara
lepidosaur lungs
couldn’t maintain multi chambered heart due to miniaturization, multichamberedness is still ontogenetically visible
axial bending, lizard
bending axis btw right/left lobes of lungs- bending to one side- one lobe reduces in volume, the other expands, air may be pumped back and forth btw, but little is moved in and out of the animal
axial bending, dog
bending axis is dorsal to thoracic cavity, sagittal ending changes thoracic volume- actively pumps air in and out of lungs for each locator cycle
sagittal plane
vertical plane which passes from anterior to posterior, dividing the body into right and left halves
7 important, independent, character developments in breathing
diaphragmatic muscles large transverse process bipedal locomotion upright posture bounding lateral stability of vertebral column endothermy
large transverse process
trunk vertebrae providing attachment sites fro axial muscles, independent of ribs; functional separation btw breathing and locomotion; characteristically large in Archosaurs
costal aspiration, reptiles
inhalation- ribs move forward and out, thorax expands, air sucked in
exhalation- ribs move backward and in, thorax compresses
craniolateral movement of ribs
forward and out
turtle breathing
use muscles
inhalation: abdominal oblique, serratus
exhalation: transverse abdominus, pectoralis
alligator breathing
craniolateral movement of ribs, have diaphragm, post hepatic septum behind liver, transversals
posthepatic septum
when pulled back, helps with breathing- ‘hepatic piston’, ‘pelvic aspiration’, muscles attached to pelvis
transversals
move liver forward- capable of breathing and walking and galloping
bird respiratory
highly modified reptilian lungs, air sacs do not exchange gases, unidirectional lungs, extract 30-35% of O2 from air, adaptation for flight, sternum moves down for inspiration; abdominal/thoracic cavities not divided (no diaphragm)
evolution of bird respiratory system
thought to be unique, findings of unidirectional flow in iguana- new understandings
mammals respiratory system
simple system, craniolateral movement, diaphragm, elastic recoil
mammal respiratory passage
mouth/nares– buccal cavity/nasal cavity– trachea– bronchi– bronchiole– alveoli– diaphragm
circulatory system involves
blood closed circulatory system (vertebrates) muscular heart arteries, veins, capillary beds portal veins
veins/arteries
veins- to lungs
arteries- from lungs
portal vein
one organ to another
simplified circulation pathway
aorta– arteries– arterioles– capillaries– venules– veins– vena cava
heart evolution
single ventricle, single aortic opening (amphibian)– single ventricle, two aortic openings (reptile)– fully divided heart (croc)
amphibian heart
oxy and deoxy blood mix in ventricle
basic heart structures
left: superioir vena cava, sinoatrial node, right atrium, inferior vena cava, tricuspid valave, right ventricle
right: left atrium, left pulmonary veins, bicuspid valve, left ventricle
middle: atrioventricular node, ventricular septum
single circulation
1V, 1A, gills, tissues, back to heart; fish, O2 picked up from gills, carried to tissue, heart doesn’t receive very oxygenated blood- possible evolutionary development of lungs
double circulation in
archosaurs, mammals, lungfish, amphibians
double circulation, single ventricle, atrium
heart– gills– air breathing organ AND tissues– back to heart from both; partly oxygenated blood coming in to lung- helps oxygenate heart; not completely oxygenated blood delivered to tissues
double and partially divided circulation, lungfish
intermediate stage; 2A, 1V– gills AND air breathing organ– from gills– tissues– back to heart; tissues receive more oxygen than non divided system
double and partially divided circulation, amphibians, reptiles
1V, 2A in middle- out right side to skin (then tissues) AND tissues– then back to heart; out left side to lung and back to heart; fully oxygenated blood going to tissues and heart
double completely divided circulation, mammals, archosaurs
2V, 2A: out RV– lung– RA– LV– tissues– RA
croc divided circulation
foramen of panizza- carries blood from LV (oxygenated) to RV to supply heart with oxygen
pulmonary
of, relating to, affecting, or occurring in the lungs; carried on by the lungs
systemic
part of the cardiovascular system which carries oxygenated blood away from the heart to the body, and returns deoxygenated blood back to the heart
coronary
pertaining to the arteries that supply the heart tissues and originate in the root of the aorta
aorta
the main trunk of the arterial system, conveying blood from the left ventricle of the heart to all of the body except the lungs
vena cava
superioir-carries deoxygenated blood from the upper half of the body to the heart’s right atrium
inferioir- carries deoxygenated blood from the lower half of the body into the right atrium of the heart
coronary support
Lamnidae: moderate- partially endothermic
Osteichthyes, Tetrapods: slight, mostly spongey
Croc, bird, mammals: extensive, no spongey (compact myocardia)
5 chamber heart
non-croc reptiles- ventricle ‘partially divided’, some division of blood but potential for mixing; blood flow can be shunted past lung to avoid build up of CO2 in lungs (diving); R-L and L-R shunts
shunt
hole or a small passage which moves, or allows movement of, fluid from one part of the body to another
croc shunt
from right atrium to foramen of panizza (R-L shunt)
euthermy
true endothermy, birds and mammals (Eutherms)
ectotherm
mainly derive body heat from external sources- radiation, conduction from the ground
endotherm
mainly derive heat metabolically, also from external environment
poikilothermy
T_b is variable (typically ectotherms)
homeothermy
single/stable T_b (typically endotherms)
temporal heterothermy
not perfectly constant homeothermy, seen in endotherms
variation in thermal physiology of vertebrates
large variation, endo/ecto and poiko/homeo grid shows organisms in all quadrants; though only mole rats are poikilo endotherms
animals in a room with homogenous T that is gradually increased
mammal- defend T_b against a gradient, maintains Tb independent of environment
snake- basically matches room T
note- real world is not thermally homogenous
real world temperature variations
snake can take advantage of microhabitats to maintain a relatively stable Tb ex. shade on a hot day, on a cold day, must be a thermoconformer
ectotherms capable of
rapid excursion of Tb- maintain ~30º, plunge into water- drop to ~10º Tb; can’t maintain optimal T during feeding
ectotherm performance curve
relative performance vs. Tb; performance increases with Tb up to a point; performance curves reach optimum around same point for all functions; Tb is tightly spread around optimum T
temperature ranges tolerable
ectotherms: -10 - 50º
active endotherm: ~30-45º
land vs aquatic ectotherms
land: ~-10 - 40º
aquatic: 5-45º
thermal conductivity, 25ºC
water: 0.58 W/m K
air: 0.024 W/m K
soil w/ organics: 0.15-2
water is 3333X air, harder for aquatic animals to reach high Tb, less steep T gradients in water- homogenized
BMR
basal metabolic rate
minimal rate of energy expenditure per unit time by warm-blooded animals at rest
metabolic rates and turpor
resting (turpor)- large drop in MR, lower Tb
resting MR < active MR, common in small birds/mammals
daily temporal heterothermy
dunnart (marsupial) drops MR below BMR and Tb decreases overnight
seasonal temporal heterothermy
ground squirrel; series of torpor events interrupted by arousal events- raise Tb to normal levels
regional heterothermy
bearded dragon lizard, under thermal stress exhibits panting behaviour to cool down (evaporation from moist inner mouth)
oral vs. cloacal T in snakes
concealed: same T
exposed: head ~10º warmer- preferentially heat head first
differential body part heating
california ground squirrel
found to have dramatic difference in T in different body parts- can elevate T of tail by allowing more blood to tail- only in response to rattlesnakes (b/c rattlesnakes have heat sensing organs)- may intimidate the snakes
emperor penguin regional heterothermy
decrease temperature of wings via wing vein, and feet, to conserve core Tb during diving
metabolic rate vs. body size
vastly different in endotherm and ectotherm
ectotherms much much lower and nearly flat line with body body mass changes
endotherms- higher metabolism for smaller animals
metabolism is CAL/GH
resting metabolic rate
SMR in ectotherms
BMR in endotherms
measured in thermoneutral zone (balancing gains and losses)
MAMR
maximum aerobic metabolic rate- higher metabolic rates when active
core temperature, heat production vs. air temperature, ectotherm
ectotherm- Tb increases with increases air T
heat production is minimal, increases w/ increasing T
core T is a passive function of air T
core temperature, heat production vs. air temperature, endotherm
at ~38ºC heat production is 3-4X larger than endotherm
decreasing air T = increasing heat production
internal T is largely independent of air T
vertebrate ectotherms
fishes, amphibians, reptiles; environment heat source, usually variable Tb, behavioural thermoreg, narrow range of ambient conditions allowing thermoreg, lower energy needs- prolonged exposure to no food, O2, larger ability to take advantage of dormancy, small sizes, long slender shapes
vertebrate endotherms
dinosaurs(?), birds, mammals, some fishes
metabolic heat source, relatively constant Tb, mainly physiological thermoreg, wide range of ambient conditions allowing thermoreg, can live in cold places, activity in cold, enhanced aerobic scope for activity
endothermic ectotherms
leatherback: largest turtle in world, globular shape, big and round, thermal inertia, can go up to Arctic circle, maintains T 8-10º, muscular activity (swimming) generate heat which is maintained, also reduce circulations to flippers
thermal inertia
maintain heat because of large size and low SA:V
regional endothermy
green turtle- active tissues ~7º warmer than water T, heat retained due to large body size and insulation of shell, increases swimming ability, facilitate long distance migration
brooding python
maintain constant T against T gradient- jacks up metabolic T by shivering, musculature activity (only during brooding)- endothermic characteristic
tunas generate heat
via red muscle; retain via counter current heat exchangers in brain, viscera, muscles
bullfish/butterfly mackerel
thermogenic organ (modified extra ocular muscle fibres, different muscles, convergence)
lamnid sharks
heat generated by slow-twitch myotomal muscle- transferred to cranial area via unique veins; contraction of extra ocular muscle also generates heat
extraocular muscle
six muscles that control movement of the eye. and one muscle that controls eyelid elevation
niche expansion hypothesis
heating of part of body- especially brain facilitates expansion in cold waters- deep diving
body temperature vs. environment
or Tb1 vs. Tb2 (different body parts)
if slope ≠ 1 some type of thermoregulation is occurring
lamnid shark and tuna convergence
hydrodynamic body form, thunniform locomotion, negatively buoyant, dive to cold depths, swim constantly with partly open mouth, similar red muscle distribution, similar tendon arrangement, endothermy (26º core), counter-current heat exchange systems; all this and not closely related
thunniform swimming
confined primarily to the caudal fin, often fin is crescent-shaped (lunate) like a small wing and connected to the body by only a thin section called the caudal peduncle
counter current exchange systems
retain generated heat
body temp vs. air temp graph
different isotherms for different populations, body adjusted to different mean temperature; homeothermic only up to certain point (30ºC in shrikes) then heterothermic
frequency and duration of turpor are a function of
feeding rate, consistant w/ idea that torpor is a body saving mechanism; negative correlation- lower frequency of torpor with high feeding rate, maintain higher Tb when well fed
chick Tb
heterothermic when born; behaviour initiated by certain minimum T’s: biting, crawling 5-10º, shivering ~15º, wing flapping ~20º, flight ~30º
endothermic performance curve
%Performance vs. Tb; can only show a narrow range of temperatures (endotherms don’t have large range of Tb)
ex. chick burst running speed- increasing, but only have data points for 30-45º
muscle performance curve (endotherms)
muscles have larger range of T (T_m), max performance is at the highest T, peak T, peak performance- looks more like a ‘traditional’ performance curve; can even plot muscle performance of endo/ectotherms together on one
why be adapted to narrow range of T?
specialists- higher peak performance than a way wider performance curve seen in a generalist 2 enzymes (1 high T acclimated, the other cold) expensive to maintain both at once, typically don't find both forms in one animal at one time
why be adapted to higher temperature range?
muscle movements create heat, body must be able to deal with high temperatures
energy requirement vs. ambient temperature
two decreasing slopes, lower one - low conductance, higher one = high conductance; both converge at same T = Tb; a specific energy requirement will cross low conductance line at lower T than the high conductance line; balance heat loss?
factors affecting conductance of a body
nature of surrounding fluid
size
shape
nature of body surface
nature of surrounding fluid (conductance)
water conductance > air
tougher to be an endotherm in water
size affects on conductance
SA:V
bigger animals lose heat less slowly
effects of shape on conductance
SA:V
rounder animals maintain heat better
nature of body surface, conductance
mammals- air, feathers, trap air btw body and surface of coat, which is a good insulator
insulating value of fur
insulating values shift with season, especially in larger mammals; having fur is not adequate to initiate endothermy- important but need the other equipment too (internal)
insulation value vs. fur thickness
positive correlation
low end- squirrels
high end- wolf, polar bear
changing conductance
if ambient temperature drops- switch from high to low conductance to conserve energy; can be done by changing erection of hairs (in fur)
generating metabolic heat
muscular contraction- physical activity, shivering
non-shivering thermogenesis
metabolism of viscera
non-shivering thermogenesis
using brown-adipose tissue, particularly well developed in young mammals
metabolism of viscera
metabolism of internal organs; 70% of heat production in mammals at rest is generated by internal organs; large internal organs in mammals
thermalneutral zone
zone of ambient T’s an animal can maintain Tb with minimal energy, complicated by conductance, BMR, and critical T’s; T_lc - T_uc
critical temperatures
T_lc lower critical- 4 possible locations: High/low BMR and high/low conductance- tightest interval is low BMR and high conductance, then high BMR high conduct., low BMR low conduct., high BMR low conduct.
high BMR, low conduct., gives widest interval on Tb but costs more E
T_lc increased by
higher Tb
higher conductance
lower BMR
TMR
torpid metabolic rate
energy expenditure vs. ambient temperature with torpor conductance
shows that even in turpor thermal neutral zone is defended
endotherm RMR, MMAR
higher resting metabolic rate- 5-10X ectotherms
higher maximum metabolic aerobic respiration
sustained activity is vastly higher, sprinting speed similar
endotherms compared to ectotherms
higher aerobic scope and endurance- more red fibres in skeletal muscle; more effective oxygen delivery; turbinate bones; erect posture, parasagittal gait; increased mitochondrial SA, leakier plasma membrane
endotherm oxygen delivery system
more vascularized lung, higher ventilation rate, 4-chambered heart; birds and mammals arrived at this independently (convergence)
endotherm mitochondria and plasma membrane
larger SA in mitochon.- higher aerobic metabolism
plasma membrane leakier to Na, K, increased action potential, muscles, osmolarity– higher activity level
turbinate bones
thin, wafer like structures, covered in moist vascularized mucosa; cool air in– past mucosa– moisture lost to mucosa– breath out– air is warmer than trniate– water is given back to mucosa– helps retain water that would otherwise be lost to environment
turnout bone problems
problems in dry environments (deserts)- potential for loss of water with each breath out
euthermy evolution
can’t tell turbinates, not preserved
bone histology can’t read much into- large ranges in endo and ectotherms
strong coronary circulation
heart well vascularized, more powerful, generate higher blood pressures, can deliver blood to distal body parts when animal is in upright position
most important character in endothermy
it is not just one character it is a whole suite of characters
advantages of mammalian euthermy
high BMR, Tb higher than Tambient, constant core Tb, high MAMR (and aerobic scope)
which advantages were the most likely targets for selection
constant core Tb, high MAMR
aerobic scope
The ratio of the maximal aerobic metabolic rate to the basal metabolic rate, typically in the range of 3–10; range of possible oxidative metabolism from rest to maximal exercise
hypotheses for evolution of endothermy
niche expansion
maintenance of high brain T
scenarios for evolution of mammalian euthermy
thermoregulation first then aerobic capacity
Aerobic capacity first
hypotheses for thermoregulation first
physiological, ecological, brain size, growth of young
hypotheses for aerobic capacity first
sustained activity, juvenile provision
most likely scenario for endothermy
correlated progression, came about by small steps; reptilians became progressively mor mammalian, gradually accumulate synapomorphies; parallel changes in different lineages (even the ones that don’t lead to mammals); integrate changes, a few at a time
stick or grip? co-evolution of adhesive topes and claw in Anolis Lizards
Crandell et al., 2014; claw characters significantly different btw arboreal and non-arboreal lizards; arboreal higher and longer, decreased curvature, wider claw tip angles; toped size and claw length/height tightly correlated
toepad
allows animal to move across smooth substrates with little difficulty (leaves, smooth bark); microscopic hair-like structures on ventral pad (setae); key innovation in anoles- niche expansion, radiation, diversification
clawed vs. non-clawed animals
clawed can occupy larger portion of habitat
claws interact with surface irregularities by
interlocking, friction
interlocking: surface irreg. > claw tip diameter
mechanical interlocking stronger, advantagous to decrease size of tip
claw curvature
highest- climbers
med- perching
lowest- ground dwelling
Nostril position in dinosaurs and other vertebrates and its significance for nasal function
Witmer, 2001; have enromous, complicated bony nasal apertures; fleshy nostril now thought to be rostral (forward); consequences for nasal air streaming, physiological parameters, circumolar odorants
nasal structure roles
olfaction, respiration, manipulation, behavioural display, thermal physiology
bony nostril
osseous nasal aperture
studies fleshy nostril using what approach
extant phylogenetic bracket
biological implications
tradition caudal position would be out of main airstream- important in convective heat loss, facilitated evaporative cooling, intermittent countercurrent heart exchange, heat and water balance, selective brain temperature regulation
true navigation in birds: from quantum physics to global migration
Holland, 2013; birds able to return to known goal from a place they’ve never been; presents conflicting findings
Type III navigational challenge
birds able to return to a goal after being displaced (even artificially) to an unknown area
true navigation
ability of an animal to return to original location after diplacement to a site in unfamiliar territory, without access to familiar landmarks, goal emanating cues, or info about the displacement route
migratory true navigation
ability of an animal to navigate to a specific breeding or wintering area following displacement
map and compass hypothesis
- determine position with respect to the goal 2. determine direction to a goal; only conducted in adult birds (experience)
celestial cues
using sun, star positions; studies find consitancy with sun compass but not sun navigation
olfactory navigation
olfactory deprivation disrupts return home; may associate odours with wind directions; lack of repeatability; without odours may use other cues
anosmia
inability to perceive odor or a lack of functioning olfaction
magnetic cues
magnetic field stronger at poles- potential to indicate latitudinal position, coarse scale; skepticism- no sense organ- earths magnetic field can prevade all tissue
radical pair mechanism
electron spin states affected by strong magnetic fields- radical pair molecule- photoreceptive- perceived through the eyes- may involve ZENK gene, cluster-N, night vision
magnetoreceptor
cryptochrome- blue light receptor, long-lived radical pairs
ferrimagnetic sense
multi doman magnetite- no magnetization, single domain- permanent magnetic moment, super paramagnetic- fluctuating magnetic moment; bacteria contain single domain, magnetite widely present in organisms; also only found in adults; magnetic field detected by trigeminal nerve
Brave new propagules: terrestrial embryos in anamniotic eggs
Martin and Carter, 2013; lots of fish and amphibians reproduce terrestrially despite absence of amniotic egg; eggs- smaller, simple chorionic membrane; disadvantage- desiccation, novel predators; arisen independently in different lineages
anamniotic egg
much smaller, more dependent on environmental conditions, simple chorion membrane
advantages of terrestrial incubation
higher T, higher O2, diffusion of O2 more rapid in air- avoid hypoxia, avoid aquatic predators
dehydrated eggs
deformed, death, hatch early
protection against dehydration, anamniotes
less aquaporin channels, amyloid fibres in the egg envelope, shape, shaded under a boulder, in a burrow, buried in damp sand
parental care, anamniotes
choice of oviposition site, guarding eggs, slashing them, exchange oxygen, transport tadpoles to water
ECH
environmental cued hatching
ECHs
decreased O2 levels, mechanical agitation of seawater, disturbance by snakes or wasps, presence of disease
influence on size of egg
yolk, offspring size, time to hatching, maternal size, habitat quality, O2 availability, duration of spawning, geographic location, parental care
endotrophy
ability to metamorphose without feeding, requires a minimum egg size
larger egg sizes provide opportunity for
developmental plasticity
types of terrestrial incubation
Type 1: conservative constitutive, Type 2: ECH early alert, 3: cautious constitutive, 4: ECH by parental involvement, 5: ECH ready and waiting, ECH ready and progressing, 7: precocious or direct development, 8: true diapauses
diapause
delay in development in response to regularly and recurring periods of adverse environmental conditions, physiological state of dormancy with very specific initiating and inhibiting conditions
propagule
any structure capable of being propagated or acting as an agent of reproduction
Developmental change in the function of movement systems: transition of the pectoral fins between respiratory and locomotor roles in zebrafish
Hale, 2014; changes in roles of morphology between stages of life history; larvae zebrafish use pectoral fins to exchange fluid near body for cutaneous respiration; musculature and positioning of fin change
developmental changes mediated by
adding cells, tissues, structures, change in body size, physics of movement, behaviour, scaling of body elements (allometry), motor control, ecological factors
fin shape
high aspect ratio (wing shape)- improved cruising performance; low aspect ratio (rounded)- high-acceleration starts and maneuvers, improved hovering stability
larval zebrafish
hatch 48-72hr post-fertilization, 3.1-3.5mm, yolk-stage -several days, pectoral fin bud forms ~24hr post-fertilization, 48h- fin elongate w/ skeleton
chaotic mixing under viscous conditions
larvae use pectoral finds pull fluid distant from the body towards the trunk and move fluid in the boundary layer away from the side of the body to increase O2
mutations in vhl gene
perceive environment as hypoxic