Final Exam Flashcards
Sagittal
Section of the body that is vertical and lengthwise to the body, dividing it into right and left parts
- Can be sagittal/midsagittal or parasagittal
Midsagittal
Sagittal section directly through the midline, dividing the body into equal halves
Parasagittal
Sagittal section parallel to the midline, dividing the body into unequal left and right sections
Transverse
Section of the body that is vertical and perpendicular to the length of the body, dividing it into front and back parts
- Perpendicular to the sagittal section
Frontal (section)
Section of the body that is horizontal and lengthwise to the body, dividing it into top and bottom parts
- Perpendicular to both sagittal and transverse planes
Anterior
Towards the head
Posterior
Towards the tail
Dorsal
Towards the back
Ventral
Towards the belly/substrate
Medial
Towards the sagittal midline
Lateral
Away from the sagittal midline
Proximal
Closer to the point of reference
Distal
Farther from the point of reference
Craniata
Largest clade within Chordata and is the clade to which the vertebrates belong
- Related to other chordates, like Urochordata and Cephalochordata
- More distantly related to Hemichordata
Pharyngotremata
Clade formed of the chordates and hemichordates
Chordata
Clade that contains the craniates, urochordates and cephalochordates
Protochordates
Group that includes everything EXCEPT the chordates
- i.e. hemichordates
- Not a natural grouping
Important Chordate Characters
- Pharyngeal slits
- Notochord
- Dorsal hollow nerve cord
- Endostyle
- Post-anal tail
Notochord
Axial structure in chordates that stiffens the midline and is flexible but resistant to compression
- Precursor to the spine
- Filled with fluid and surrounded by collagenous and connective tissue sheaths
Hemichordata
Sister group to Chordata
Two clades
- Enteropneusta
- Pterobranchia
Pterobranchia
Clade within the Hemichordata
- Filter-feeder or suspension feeder
- To feed, secretes a sticky fluid onto a proboscis that projects just beyond the proboscis/mouth
- Lopophores (tentacles) help move food to mouth
- Live in tube-like structures when living in colonies
Lacks chordate characters:
- No post-anal tail
- No pharyngeal slits (most of the time)
- No dorsal hollow nerve cord
- No endostyle (most of the time)
- No notochord
Enteropneusta
Acorn worms
Clade within the Hemichordata
- Filter feeder that waits for prey to fall into its burrow
- Has many pharyngeal slits (used for water expulsion)
- Has dorsal nerve cord
Lacks chordate characters:
- No post-anal tail
- No endostyle
- No notochord
Endostyle
Mid-ventral groove that secretes mucus
Urochordata
Tunicates
Clade within Chordata
- Has a large, barrel-shaped pharynx with many pharyngeal slits
- Has an endostyle
Characters in the juvenile
- Post-anal tail with a notochord and dorsal nerve cord
- Endostyle and pharyngeal slits, found in adult, are both present as well
- All five chordate characteristics ONLY present in the juvenile
Cephalochordata
Branchiostoma/amphioxus
Clade within Chordata, most closely related to craniates
- Have all typical chordate characters
- Notochord extends into the head
- Pharyngeal slits empty into a common chamber
- Filter feeders that sit buried in the substrate except for their head
- Poor swimmers despite fish-like body (lack paired fins)
- Muscular bundles segmented along the body wall
- Has no true brain and poorly developed sense organs
Heterochrony
A relative change in developmental timing that leads to changes in size and/or shape
Paedomorphosis
The retention of juvenile features in the adult of a descendent
- Adult retains characters present in the young of the ancestral stage
- Can do this in many ways: slow somatic development, speed up sexual development
Implication for vertebrates:
- The larval stage of urochordates have all chordate characteristics
- Higher chordates may have developed from an ancestor that initially resembled the larval urochordate
Craniata
Clade within Chordata
- Refers to chordates that have a well-defined “head” and a hard skull
Agnathans
Craniates that lack jaws
- Also lack paired appendages
- Only extant species are the cyclostomes, used to also include ostracoderms
Ostracoderms
Extinct, armoured agnathans
- Covered by bony plates
- Jawless
Cyclostomes
Agnathans that include the Petromyzontoidea and Myxinoidea
- Parasitic, rasping tongue allows them to burrow into prey
- Entirely cartilaginous skeleton
- Single median nostril
- No true teeth - keratinous “teeth” known as denticles
Petromyzontoidea
Lampreys
- Predatory, parasitic
- Oral cup attaches to prey, allowing them to drink blood and other body fluids
Myxinoidea
Hagfish
- Scavengers
- Use horny tongue to rasp flesh from dead or dying prey
Denticles
The keratinous teeth of cyclostomes
Gnathostomata
True, jawed fishes
- Includes the placoderms and eugnathostomes
Placodermi
Extinct gnathostomes
- Usually have lots of bony armour
- Have jaw-like structures, but these are made from their armour (dermal bone) rather than branchial arches (visceral bone/cartilage)
- Paired fins
Eugnathostomata
“True jawed vertebrates”
- Jaw made from visceral structures, either bone or cartilage
Two major groups:
- Chondrichthyes
- Teleostomi
Chondrichthyes
Sharks and relatives Basal eugnathostomes - Cartilaginous skeleton with very few bits of bone (secondary loss) - Well-developed, paired fins - Two nostrils - Males usually have a clasper
Two main clades:
- Elasmobranchii
- Holocephalia
Elasmobranchii
Sharks, skates and rays
- More diverse than their sister group, the holocephalians
- Upper jaw is movable since it is free from the braincase
- Gills open directly to the environment
- Rapid tooth replacement
- Most species have an elongated, fusiform body
Holocephalia
Chimaeras and ratfish
- Much less diverse than their sister group
- Slow tooth replacement
- Pharyngeal slits are covered by an operculum (flap of skin)
- Upper jaw is usually fused to the braincase
- Tend to be benthic mollusc feeders, so teeth are flattened to be used in grinding
Stages leading to the typical vertebrate condition
- Prevertebrate stage
- Agnathan stage
- Gnathostome stage
Prevertebrate Stage
Urochordates, cephalochordates
- Suspension feeders (cilia-mucus)
- Small, poor swimmers
- Ciliary action produces movement of water
Agnathan Stage
Agnathans (durrr)
- Muscular bands plus cartilaginous bars in the pharynx produce a muscular pump to move water for feeding
- Still a filter feeder, but on larger particles due to larger size
- Absence of paired fins make them poor swimmers
Gnathostome Stage
Jawed vertebrates
- Development of jaws allows selection of larger prey and hunting
- Larger prey allows for larger size
- Problem of how to achieve gas exchange solves through addition of gills to pharyngeal slits
Higher Fishes
Term that is usually used to refer to Osteichthyes
Osteichthyes
“Bony fishes”
- Bony skeleton is retained and enhanced
- Most successful vertebrates in terms of diversity and numbers
Groups:
- Actinopterygii
- Sarcopterygii
Teleostomi
Clade that includes Acanthodii and Osteichthyes
Acanthodii
Known as "spiny sharks" (not sharks) Extinct - Heterocercal tail - Bony plates as well as scales - Have characteristically strange paired fins that are supported by a spine; more than two pairs
Actinopterygii
Ray-finned fishes
- Most successful of the bony fishes (99% of extant species)
- Inhabit all marine and freshwater habitats
- Fins supported by rays, so have little muscular development
- Primitively had one dorsal fin and ganoid scales
- Isolated nasal sacs with no internal nostril
Three types
- Chondrostei
- Holostei
- Teleostei
Sarcopterygii
Lobe-finned/Fleshy-finned fishes
- Few extant species, but used to be more successful
- Ancestors of terrestrial vertebrates… so all tetrapods are sarcopterygians
- Fins have well-developed skeletal support and strong muscular development
- Have cosmoid scales and two dorsal fins
- Nasal opening extends into the oral cavity
- Diphycercal tail
Extant species
- Dipnoi
- Coelecanth
Dipnoi
Lungfishes
Coelecanth
“Latimeria”
Chondrostei
Early, primitive actinopterygians
- Few extant species, inc. sturgeons
- Hyomandibular slopes backward
- Long jaw
- Maxilla involved in biting
- Heterocercal tail
Holostei
Not a natural group Intermediate actinopterygians - Freshwater species that are typically fast-swimming - Jaws shortened - Hyomandibular more or less vertical - Premaxilla & maxilla tightly fused - Premaxilla elongated - Reduced heterocercal tail, more symmetry
Teleostei
Most living fishes
- Great diversity in body shape and activity
- Hyomandibular slopes forward
- Premaxilla expands to free the maxilla and exclude it from biting
- Maxilla allows premaxilla to slide back and forth
- Jaws can be protruded when mouth opened, allowing for greater control and different kinds of biting
Trends in Actinopterygian Body shape and fin form
Early
- Fusiform body shape
- Fast-swimming
- Fins positioned for optimal gliding
Advanced
- Body more shortened
- Pectoral fins move dorsally, to be used as brakes and allow for greater control
- Pectoral fins move anteriorly
Heterocercal
Tail in which the posterior end of the body curves upward so that there are more fin rays ventrally on the tail
Homocercal
Caudal fin is symmetrical superficially, but internally the vertebrae still curve dorsally
Diphycercal
Tail type in which the caudal fin is symmetrical both superficially and internally
Tetrapoda
Vertebrates whose ancestors made the transition to terrestrial life
- Breathe air
- Are mainly amphibious or terrestrial, although some specialized descendants returned to the water
Two main groups
- Amphibia
- Amniota
Stegocephalia
Group that includes the tetrapods and their non-terrestrial, but still limbed, ancestors
- Fully-formed limbs, skull and vertebrae able to support weight on land
- Basal species include Icthyostega and Acanthostega, which were not well-adapted to life on land
- Still retain branchial arches and gills, so were most likely aquatic
Elpistostegids
Sarcopterygians that have made some changes to be similar to tetrapods but are still clearly fish
- Have internal nostrils
- Includes Tiktaalik, who is closely related to stegocephalians
Amphibia
Biologically intermediate between fishes and more derived tetrapods
- In general, lay eggs in water and have an aquatic larval stage; this means they are restricted to a moist environment
Living group: Lissamphibia
Amniota
More derived tetrapods
- Lay amniotic eggs
Two major lineages
- Reptilia
- Synapsida
Amniotic Egg
Egg type in which extra-embryonic membranes surround and protect the developing embryo
- Prevents desiccation
- Engages in gas exchange
- Results in a longer developmental period
- Young emerge as essentially miniature adults, no metamorphosis
Lissamphibia
Extant amphibians
- Some still retain ancient forms of locomotion, but are notably not primitive
- Cutaneous respiration is the primary form of gas exchange
Three groups
- Caudata (salamanders)
- Anura (frogs)
- Gymnophiona (caecilians)
Anapsid
Skull type in which there are no temporal fenestrae
- Typically present in basal amniotes, but debate over if its an ancestral form
- Still retained in turtles
Euryapsid
Skull type in which there is a single temporal fenestra (supratemporal)
- Seems to be descended from diapsid skull through secondary closure of the infraorbital fenestra
Diapsid
Skull type in which there are two temporal fenestrae
- Most common form
Synapsid
Skull type in which there is a single temporal fenestra (infratemporal)
Reptilia
Diverse lineage of tetrapods including lizards, birds, dinosaurs, etc.
Two main groups
- Parareptilia
- Diapsida
Parareptilia
Reptiles with anapsid skulls
- Inc. Testudines
Testudines
Turtles
Diapsida
Large, diverse group of reptiles
- Diapsid and euryapsid skull
- Diverse even by standards of all of Tetrapoda
Two main groups
- Archosauromorphia
- Lepidosauromorphia
Archosauromorphia
Includes:
- Crocodiles
- Pterosaurs
- Dinosaurs (inc. birds)
Lepidosauromorphia
Includes:
- Ichthyosaurs
- Plesiosaurs
- Rhynchocephalians
- Lizards (inc. snakes)
Synapsida
One of the major clades within Amniota
- Includes Mammalia and their fossil relatives
- Characterized by synapsid skull type
Two broad groups
- Pelycosaurs
- Therapsida
Pelycosaurs
Early, basal group of synapsids
- Not a natural group
Therpasida
More derived group of synapsids
- Includes mammals
- Some early species had intermediate leg positioning, with hind limbs under body but forelimbs swung out to the side
- Several groups in addition to mammals, including symmetrodonts, multituberculates and triconodonts
Mammal-like Reptiles
Term used to refer to all non-mammalian synapsids
- Reflects the idea that they were more reptile-like in their general way of living
- Transitional stages towards mammalian condition have all four bones (squamosal, quadrate, articular, dentary) involved in jaw bones
Mammalia
Group of therapsids that represents all mammals
Characterized by:
- Mammary glands
- Hair
- Muscular diaphragm
- External ears (a few exceptions)
- Mandibular halves formed by single dentary
- Squamosal/dentary jaw joint
- Three middle ear ossicles: malleus, incus, stapes
- Teeth tend to be complex with precise occlusion
- Body held upright instead of sprawling
Two groups
- Monotremata
- Theria (inc. Marsupialia & Eutheria)
Relation between ear ossicles and jaw joint in mammals vs. reptiles
The two additional ossicles in mammals are derived from the mandibular bones in reptiles
- Articular = malleus
- Quadrate = incus
- Stapes = stapes
In addition, the angular in the reptilian jaw forms the ectotympanic in mammals
Tribosphenic Molar
Form of molars where the upper molars have three cusps and the lower molars have four
Bunodont Molar
Form of molars where both the upper and lower teeth have four cusps
Monotremata
Group of mammals that includes only three extant species
- Platypus (Ornithorhynchidae) & 2 sp. of echidna (Tachyglossidae)
- Have hair, suckle young and are endothermic
Ancestral features
- Cloaca
- Absence of nipples
- Lay eggs in a leathery shell
Marsupialia
A.K.A. Metatheria
- Much more diverse group than monotremes, but less so than placental mammals
- Found in Australia and South America (as well as their surrounding regions)
- Females of many species have a pouch in which they carry their young
- Young are usually born at an early stage and complete their development in their mother’s pouch
- Have a placenta, but it is a yolk sac placenta and is not vascularized except in peramelids
Eutheria
A.K.A. placental mammals
- Much more numerous, diverse and widespread than marsupials and monotremes
- All species have a chorioallantoic placenta
- Young remain in utero until they are developmentally more advanced before being born
- Occupy many niches, including the air, water and land
Chorioallantoic Placenta
Type of placenta that is vascularized
- Found in peramelids and eutherians
- Comes at a price: can’t dispose of offspring that are too energetically costly
Embryology
The processes involved in early development of a fetus
- Important process for the animal
- Characters can be used to study phylogenetic relationships
- Understanding helps us make sense of the organization of various systems
Early embryological processes
Vary greatly among vertebrates but can be generally simplified into three or four patterns
- All lead to more or less the same end: gastrula with similarly arranged primary body tissues
- Further differentiation can be more easily compared
Microlecithal
Egg type in which there is little yolk
- E.g. in amphioxus
- Blastula is one cell thick
Mesolecithal
Egg type in which there is a moderate amount of yolk
- Initial divisions less equal than in microlecithal egg
- Cell of the animal pole are smaller than in microlecithal
- Blastula is several cells thick
- Blastocoele is displaced towards the animal pole
Macrolecithal
Egg type in which there is a lot of yolk
- Essentially a big mass of yolk with a small amount of protoplasm on top
- Only small cap of protoplasm divides, so it is difficult to compare to division in other egg types
- Blastula is a plate of cells at the animal pole that is separated from the yolk by the blastocoele
Yolk
Food material that is inert to an egg
Vegetal Pole
End of an egg towards which the yolk gravitates
- Lower hemisphere of the egg
Animal Pole
End of an egg that is less yolky
Early divisions of an egg cell
Occurs in the protoplasm of an egg
- Vertical
- Vertical
- Horizontal
Blastula
Spherical form of an egg after continued divisions of the protoplasm
- Actually a plate of cells in macrolecithal eggs
Blastocoele
Central cavity found within a blastula
- Separates blastula from yolk in macrolecithal eggs
Mammalian Embryo Development
Different from egg forms, since the embryo develops in the uterus
- Spherical mass is formed like in microlecithal eggs but the resemblance is only superficial
- Becomes a trophoblast with an inner cell mass
Trophoblast
Spherical mass of cells that forms in mammals after the continued division of the protoplasm
- Has an inner cell mass toward the animal pole
Inner Cell Mass
Cluster of cells towards the animal pole within a trophoblast
- Later forms the embryo and the extra-embryonic membranes
Primary Germ Layers
Basic arrangement of the body
- Become delimited during gastrulation then become differentiated into various body regions
- End up producing all the major body organs
- Each layer produces the same structures in adult vertebrate bodies, with few exceptions
Three layers
- Ectoderm
- Mesoderm
- Endoderm
Gastrula
Forms through continued cell division, migration and differentiation in the blastula
- Vegetal pole folds into a cup-like structure, mainly by the rolling of cells into the blastocoele
- Cells of the animal pole form the outer layer while the vegetal pole forms the lining of the gut
Gastrocoele
A.K.A. Archenteron
Central cavity within a gastrula
- Formed from pocketing of of the blastula as it forms the gastrula
Blastopore
Entrance into the gastrocoele
Ectoderm
Germ layer formed from the animal pole of an egg
- Becomes the skin and nervous system
Also includes: neurectoderm
Mesoderm
Germ layer between the ectoderm and endoderm
- Becomes the somatic skeleton, muscle and circulatory system
Two kinds
- Chordamesoderm
- Lateral mesoderm
Endoderm
Germ layer formed from the vegetal pole of an egg
- Becomes the digestive tract, visceral muscles and visceral skeleton
Neurectoderm
Forms from the dorsal middle of the ectoderm
Chordamesoderm
Portion of the mesoderm that becomes the notochord
Lateral Mesoderm
Portion of the mesoderm that forms pretty much everything but the notochord
Development of the Ectoderm
- Partially differentiates into the neurectoderm dorsally
- Neurectoderm inpockets and ectoderm expands to cover it
- Neural crest cells and neurogenic placodes form
- Ectoderm covers the neural tube
- Neural crest cells begin to migrate
Development of the Mesoderm
- Differentiates into the chordamesoderm and lateral mesoderm
- Mesoderm becomes segmented into somites and expands laterodistally
- Lateral mesoderm subdivides, forming the coelom
- Completely comes between the ectoderm and endoderm
- Lateral mesoderm subdivides into the epimere, mesomere and hypomere as the coelom enlarges
Development of the Endoderm
- Expands dorsomedially
- Completely surrounds the gastrocoele
- Becomes relatively smaller
Neural Crest Cells
**In notes
Neurogenic Placodes
**In notes
Epimere
Dorsal subdivision of the lateral mesoderm
Mesomere
A.K.A. nephrotome
Middle subdivision of the lateral mesoderm
- Becomes the kidney, excretory ducts and reproductive ducts
Hypomere
Ventral subdivision of the lateral mesoderm
Development of the Epimere
- Subdivides lateromedially into the dermatome, myotome and sclerotome
Dermatome
Lateral subdivision of the epimere
Myotome
Middle subdivision of the epimere
Sclerotome
Medial subdivision of the epimere
Development of the Hypomere
- Coelom expands further
- Expands dorsomedially to enclose the endoderm
- Subdivides into a somatic layer and a visceral layer
Development of the Dermatome
- Expands deep to the ectoderm
2. Loses segmentation
Development of the Myotome
- Expands dorsally between the ectoderm and sclerotome and ventrally between the ectoderm and hypomere
- Is subdivided by a horizontal skeletogenous septum
Development of the Sclerotome
- Surrounds the notochord and the neural tube
Structures formed from the Ectoderm
- Epidermis and derivatives
- Lens of the eye
- Anterior lining of the oral cavity
- Various sensory head structures (neurogenic placodes)
- Visceral skeleton, including jaws
- Meninges
- Structures of the nervous system
- Neural tube: brain, spinal cord, nerves
Structures formed from the Dermatome
- Dermis of the integument
- Dermal muscles
- Dermal skeletal structures
Structures formed from the Myotome
- Skeletal muscles: epaxial and hypaxial
- Appendicular skeleton
Structures formed from the Sclerotome
Vertebral column
Structures formed from the Visceral Hypomere
- Visceral serosa, including mesenteries
- Visceral musculature
- Heart
- Blood vessels
- Lymph vessels
- Gonads
Structures formed from the Somatic Hypomere
Parietal serosa
Structures formed from the Endoderm
- Liver
- Pancreas
- Lining of the lungs
- Digestive tract
- Urinary bladder
Integument
A.K.A. Skin
Acts as the covering for the body and is one of the largest organs
- Protects the body from UV radiation, mechanical forces, micro-organismal invasion
- Acts as camouflage and colouring
- Regulates temperature
- Creates claws, hair, feathers, nails, horns, etc.
- Can be used for respiration
- Tends to become thicker as species become more derived, but many vertebrates still have (semi)permeable skin
Primary Body Tissues
- Epithelial
- Connective
- Muscular
- Nervous
Epithelial Tissue
Layer(s) of cells that forms a barrier
- Covers exposed surfaces as part of the skin
- Lines body cavities
- Cells are tightly bound together
- Usually avascular
- Have few, if any, nervous structure
- Has apical, exposed surface and a basal surface that attaches to the basement membrane that connects it to underlying body structures
Serosa
Epithelial tissue that is internal and has no exit
- Thoracic: pleura
- Abdominal: peritoneum
- Blood vessels: endothelium
Mucosa
Epithelial tissue in structures that have an exit
- Respiratory tract
- Digestive tract
Epidermis
Superficial, epithelial layer of integument
- Formed from the ectoderm
- Usually thin
- Avascular
- Has almost no nervous structures
Two main layers (five total)
- Stratum corneum
- Stratum basale
Dermis
Deep, connective tissue layer of integument
- Formed mainly from the dermatome
- Usually relatively thick
- Vascularized
- Has nervous structures in the form of sensory receptors
Issues with (Semi)Permeable Skin
When away from water, the organism will lose internal water quickly
- Even marine vertebrates lose water because they are less than 50% seawater
“Water-Proofing” Skin
Change the nature of the epidermis so that the most superficial parts become differentiated into a layer of dead, hardened cells
- Incorporates insoluble proteins like keratins
- Skin considered cornified or horny
Hypodermis
A.K.A. Superficial fascia
Deep to the dermis
- Mainly loose connective tissue and adipose connective tissue
Stratum Corneum
Superficial layer of the epidermis formed from dead cells
- Thin
- Formed of dead flattened keratin-filled keratinocytes that have been pushed up from the stratum basale
Stratum Basale
A.K.A. Germinativum
Deep, living, layer of the epidermis that contains dividing cells
- Most cells are keratinocytes, which are constantly being formed there
- Keratinocytes pushed superficially as new ones form and take on keratins
Integument in Fishes
Skin is relatively thin
- Epidermis has few keratinized cells, so is mostly living
- Secretes mucus that helps protect against infectious bacteria, makes the fish slippery, helps reduce drag and contains various chemicals
- Sometimes forms specialized keratinized structures, such as denticles in the lamprey
Integument in Derived Fishes
Scales usually present
- Scales formed from the epidermis and dermis
- May include enamel (epidermal), dentin (dermal) and bone (dermal)
Placoid Scales
Type of scale seen in sharks
- Scale is formed from dentin and projects through the epidermis
- Capped by enamel
- No dermal bone
Dermal Bone in Fish
- Forms the bony plates on the head and trunk in ostracoderms and placoderms
- More posteriorly, bone is broken into smaller dermal scales; also present over actinopterygians and sarcopterygians
Bony Fish Scales
Formed by dermal bone and may be capped by enamel and dentin
Different Types
- Cosmoid Scales (primitive sarcopterygians)
- Ganoid Scales (primitive actinopterygians)
- Teleost Scales (derived actinopterygians): Cycloid or Ctenoid
Integument in Amphibians
Tends to be thin and composed entirely of live cells
- Stratum corneum is thin and provides minimal protection against mechanical abrasion and water loss
- Scales are absent except in some caecilians
- Cutaneous respiration is extremely important, so capillaries reach into the lower epidermis
- Generally have mucus and poison glands
Integument in Reptiles
Skin that has more extensive keratinization, with scales
- Dermal bone may be present but is not usually associated with scales
- Osteoderms and gastralia may be present
- Few skin glands that are mainly scent glands
Reptilian Scales
- Usually lack underlying bony dermal contribution
- Form from a fold in the surface epidermis
- Can be modified into crests, spines and “horns”
Osteoderm
Small piece of dermal bone found under epidermal scales
- Found in some turtle shell bones, crocodilians, some lizards and extinct reptiles
Gastralia
Collection of dermal bone in the abdominal region
Epidermal Derivatives
- Scales
- Calluses
- Nails
- Claws
- Hooves
- Hair
- Feathers
- Baleen
- Horns
- Antlers
- Glands
Nails, Claws and Hooves
Keratinized epidermal structures that tip the digits of amniotes
- Protective
- Functional in climbing, defence, food gathering, etc.
Horns and Antlers
Widespread cranial appendages that are used in defence, mate selection and dominance
- True horns are a keratinized sheath supported by an unbranched bony core/spike; neither are shed
- Antlers are usually confined to males and are a velvety shin until becoming bone when mature; shed annually; usually branched
- Rhino horn is keratinized epithelium in the form of fused, hairlike epidermal papillae
- Giraffe horns are ossified cartilage cores covered by skin
Hair
An exclusively mammalian feature that is formed from an epidermal sheath (follicle) that extends into the dermis
- Grows basally with dermal papilla supplying blood vessels
- Cells are pushed up, keratinized and die
Arrector Pili
Muscle associated with a hair follicle
Sebaceous Gland
Gland associated with a hair follicle that secretes an oily substance to lubricate and protect the hair
Apocrine Gland
Sweat glands that are associated with hair follicles in restricted areas
- Secrete viscous substances or pheromones
Glands
Epidermal structures found in all vertebrate clades Can be single celled or multicellular: - Alveolar - Tubular/coiled - Complex or compound
Uropygial Gland
Gland near the base of the tail in birds that secretes an oily substance used in preening
- Protects feathers and repels water
Salt Gland
Gland on the head of some birds that is used to secrete excess salts
Eccrine Gland
Gland that secretes salt, urea or water
Mammary Gland
Modified apocrine gland used to secrete milk
Scent Gland
Modified apocrine gland used to mark territory, recognize other individuals and courtship
Skeleton
Supportive structure that is composed of various kinds of connective tissue
Connective Tissue
Consists of cells scattered through a matrix that is normally secreted by the cells
- Formed from mesenchyme
- Reinforces the epithelia and other soft body tissues
- Acts as a support for the body
- Proportion of cell types and materials involved determines structure, properties and function
Mesenchyme
Embryonic material that is formed mainly from the epimere and hypomere
- Forms a network of cells between the outer tube and developing organs
In adult, differentiates into
- Connective tissue
- Most of the circulatory system
- Most of the muscles
Bone
Connective tissue of cells (osteocytes) deposited in a heavily mineralized matrix
- Extremely metabolically active due to high vascularization
- Osteocytes reside in lacunae within the matrix; connect with others
- Grows by expansion at the surface
Cartilage
Connective tissue of variable types
- Matrix is normally rigid but not as hard as bone
- Chondrocytes reside in spherical, isolated lacunae
- Not very metabolically active due to being avascular
- Grows through both internal expansion and expansion at the surface
Bone vs Cartilage
It used to be thought that cartilage was primitive to bone, an idea that seemed to have embryological support since cartilage forms before bone in the embryo
- However, the most primitive vertebrate so far recovered had an extensive bony exoskeleton and since then bone has been lost secondarily
- Now considered to simply be different kinds of skeletal tissue
- Bone better for compression
- Fibrous cartilage better in tension
Parts of the Skeleton
Components that vary in structure, position and embryologic origin
Three broad classes
- Dermal skeleton
- Endoskeleton
- Heterotopic skeletal elements
Dermal Skeleton
Membranous bone that forms directly in connective tissue
- Relatively superficial in position
- Forms mostly in the dermis
- Never preformed in cartilage
Examples
- Bony plates/scales in early fish
- Osteoderms in crocodilians/turtles
- Parts of the skull
Endoskeleton
Deeper lying bone that is almost always endochondral in formation
- Replacement of embryonic cartilage by adult bony structures
- Forms most bones in the body
Subdivisions
- Somatic Skeleton
- Visceral Skeleton
Somatic Skeleton
Skeleton from the outer tube of the body that is formed by somites
Two types:
- Axial
- Appendicular
Axial Skeleton
Most of the skull, vertebral column and axial ribs
Appendicular Skeleton
Paired appendages and limb girdles
Visceral Skeleton
Skeleton that is associated with branchial structures or derived from them
- Formed by the neural crest cells
Heterotopic Skeletal Elements
Stray skeletal elements that are not associated with other parts of the skeleton
- Not usually preformed in cartilage
Examples
- Sesamoid bones
- Bone in diaphragm in camels
- Os cordis: bone in septum of heart in some ruminants
- Bone in upper eyelid in crocodilians
- Baculum, os clitoris
Diaphysis
Shaft of a long bone
Epiphysis
End of a long bone
Epiphyseal Line
Line that separates the diaphysis and epiphysis
- Formed from fusion of the epiphyseal plate
Epiphyseal Plate
Layer of hyaline cartilage that is found in juveniles between the epiphysis and diaphysis
Periosteum
Fibrous connective tissue that surrounds a bone
- Has a deep, bone-forming layer
Medullary Cavity
Cavity within the diaphysis of a bone within which blood cells are formed
Endosteum
Connective tissue lining the medullary cavity and spaces within a bone
Articular Cartilage
Cartilage found at the joints of bones
Development of Endochondral Bone
- Begins as a model of hyaline cartilage
- Bone begins to grow, radiating from the centre as the bone grows in length and the medullary cavity begins
- New centres of bone growth begin at the ends of the bone (epiphyses)
- Bone grows in width and new bone forms at the epiphyseal plate
- Epiphyses fuse and bone ceases growth
Postcranial Skeleton
All skeletal structures of the body not including the head
- Largely formed by somatic endoskeletal elements
- Axial and appendicular
- Mesodermal in embryological origin
Vertebral Column
Major midline axial structure element
- Formed by a series of repeating skeletal elements along the back
- Little musculature attaching directly to vertebrae in fish, with most muscular force exerted on myosepta
- Thus, ribs develop at the intersection of connective tissues to make muscular effort more efficient
- Type of rib depends on position
Dorsal Rib
Typical rib that occurs at the intersection of the myoseptum and horizontal skeletogenous septum
- Formed from endochondral bone
Ventral Rib
Ribs that occur in fishes at the intersection of the myoseptum and connective tissue along the coelomic cavity
- Not to be confused with gastralia and abdominal ribs
- Formed from endochondral bone
Vertebrae
Each formed by various parts
- Centrum is main structural element and articulates at either end with other centra
- Different types depending on the form of their ends
Types
- Amphicoelous
- Procoelous
- Opisthocoelous
- Acoelous
- Heterocoelous
Amphicoelous
Type of vertebra where the centrum is concave at both ends
- Typical of fishes and early tetrapods
Procoelous
Type of vertebra where the centrum is concave anteriorly and convex posteriorly
- Typical of amphibians and reptiles
Opithocoelous
Type of vertebra where the centrum is convex anteriorly and concave posteriorly
- Typical of amphibians and reptiles
Acoelous
Type of vertebra where the centrum is nearly flat on both ends
- Typical of the trunk of mammals and birds
Heterocoelous
Type of vertebra where the centrum is saddle shaped
- Typical of the neck vertebrae of birds
Evolution of Vertebrae
Tendency for there to be increased differentiation along the column in more advanced forms
- Fish have trunk and caudal
- Tetrapods have trunk, caudal and one or more sacral
- More advanced tetrapods have cervical, dorsal, sacral and caudal
- Mammals have cervical, thoracic, lumbar, sacral and caudal
Embryologic Development of Vertebrae
Develop from the sclerotome and are initially segmentally arranged
- In fish, can be segmentally or intersegmentally arranged
- In amniotes, become intersegmentally arranged with respect to musculature
- Ensures musculature forms across adjacent vertebrae, otherwise muscle would not have functional role
- Primary sclerotome migrates medially and toward the notochord and become rearranged
- Form secondary sclerotome blocks made of cells that are originally from two adjacent primary sclerotomes
Appendicular Skeleton of Fish vs Tetrapods
Fish
- Relatively simple and small
- Pectoral > Pelvic
Tetrapods
- Large and more complex; needed to keep body off ground and propel it
- Pelvic > Pectoral
Pectoral Girdle of Bony Fish
Have lots of dermal bone in the skin
- Pectoral girdle formed from dermal and endoskeletal bone
- Fin supported by endoskeletal bone
- Dermal elements add strength and anchor the pectoral girdle to body, mainly behind operculum; clavicle ventrally and dorsally attaching girdle to skull
- Endoskeletal deep and posterior to dermal
Pectoral Girdle of Tetrapods
Gills have been lost, so a moveable neck is possible
- Dermal elements are retained from fish except the dermal series
- Cleithrum and clavicle strongly reduced
- New interclavicle lies between the clavicles
- Both cleithrum and interclavicle lost in more advanced vertebrates
- Endoskeletal bones expand and the scapula begins to dominate
- In mammals, essentially only the clavicle remains of the dermal elements, which tends to be reduced in cursorial mammals allowing for a more mobile scapula
Pelvic Girdle of Fish
Relatively small and is not attached to the body/vertebrae
Pelvic Girdle of Tetrapods
Fused to one or more sacral vertebrae through a dorsal extension of the girdle
- Hind limbs provide nearly all locomotive force
Evolution of Posture in Tetrapods
Early Tetrapods
- Limbs perpendicular to body axis and extend laterally, causing a sprawled posture
- Locomote mainly by fish-like movements
First Change: manus and pes rotate to point forward, but lateral bending of the body still propelled them
Later Tetrapods (Archosaurs, Therapsids)
- Limbs tucked underneath body, more aligned to parasagittal plane
- Appendages use for-aft motion to create more efficient locomotion
Skull
The skeletal structure of the head minus the lower jaw
- Complex fusion of dermal and endoskeletal elements
Shark Skull
Consists of a chondrocranium and palatoquadrate
- Missing the dermal skull elements
- A single structure that begins as separate cartilaginous centres of formation
- Occipital region posteriorly: surrounds foramen magnum and occiptal condyle
- Otic capsules: anterior widening of occipital region
- Orbital region: narrowing anterior to the otic capsules
- Ethmoid region contains nasal capsules
- Braincase articulates ventrally with palatoquadrate
Braincase
The box surrounding the brain and sense organs to support and protect them
- Complex series of single and paired bones
- Mostly somatic endochondral bone
- Some dermal bone
Dermal Bone
- Parasphenoid
Somatic Endochondral
- Supraoccipital
- Basioccipital
- Exoccipitals
- Opisthotic
- Prootic
- Basisphenoid
- Sphenethmoid
Vertebrate (Bony) Skull
Three main parts
- Braincase
- Palatal complex
- Skull roof
Palatal Complex
Series of bones that comprise the palate and articulation with the lower jaw
- Has major articulations with other skull structures: lower jaw (via quadrate) and braincase (at basisphenoid via the epipterygoid)
- Paired bones that are fairly complex
- Mostly dermal bone
- Some visceral endochondral bone
Dermal Bones (bear teeth in fishes and early tetrapods)
- Pterygoids
- Ectopterygoids
- Palatines
- Vomers
Visceral Endochondral Bones
- Palatoquadrate
- Quadrate
- Epipterygoid
Skull Roof
Acts as a dorsal shield for the skull in bony vertebrates
- Nearly solid with openings for the nostrils, eyes and pineal eye
- Primitively notched posteriorly in the ear region
- Dermal bone, generally paired
Five main series of bone
- Tooth-bearing, marginal series
- Midline series
- Circumorbital series
- Temporal series
- Cheek series
Tooth-bearing Marginal Series
Series of the skull roof that forms the outer rim
Includes
- Premaxilla
- Maxilla
Midline Series
Series of the skull roof with adjacently paired bones on the dorsal side
- Nasal
- Frontal
- Parietal
- Postparietal
Circumorbital Series
Series of the skull roof that surrounds the orbit
- Prefrontal
- Postfrontal
- Lacrimal
- Postorbital
- Jugal
Temporal Series
Series of the skull roof that lies between the otic notch and the midline series
- Intertemporal
- Supratemporal
- Tabular (?)
Cheek Series
Series of the skull roof that lies behind the orbit and below the otic notch
- Squamosal
- Quadratojugal
- Quadrate
Pterygoid
Largest bone of the palatal complex Paired - Largest of the four - Posterior - Median - Dermal bone
Ectopterygoid
Bone of the palatal complex
Paired
- Posterior and lateral
- Dermal bone
Palatine
Bone of the palatal complex
Paired
- Lateral and between the ectopterygoid and vomer
- Dermal bone
Vomer
Bone of the palatal complex
Paired
- Anterior and lateral
- Dermal bone
Palatoquadrate
Bone of the palatal complex
Paired
- Visceral endochondral bone
- Largely replaced functionally by dermal elements, but usually remains as a reduced structure
- Four ossifications may form along it in fishes, but only two in tetrapods: quadrate and epipterygoid
Quadrate
Bone of the palatal complex Paired - Ossification of the palatoquadrate in tetrapods - Small and posterior - Articulates with the lower jaw - Visceral endochondral bone
Epipterygoid
Bone of the palatal complex Paired - Ossification of the palatoquadrate in tetrapods - Large and anterior - Articulates with the braincase
Parasphenoid
Bone of the braincase
Paired
- Forms the ventral part of the braincase
- Narrow anteriorly and wide posteriorly
- Dermal bone
- Forms from the skin on the roof of the oral cavity
Supraoccipital
Bone of the braincase that forms part of the occipital
Single
- Dorsal to the foramen magnum
Basioccipital
Bone of the braincase that forms part of the occipital
Single
- Ventral to the foramen magnum
Exoccipital
Bone of the braincase that forms part of the occipital
Paired
- To the left and right of the foramen magnum
Occipital
Single bone that forms the posterior part of the skull and is typically fused from the supraoccipital, basioccipital and exoccipitals
Opisthotic
Bone of the braincase that is associated with the otic region
Paired
- Posterior to the prootic
- Contains the sacs and canals of the inner ear
Prootic
Bone of the braincase that is associated with the otic region
Paired
- Anterior to the opisthotic
- Contains the sacs and canals of the inner ear
Basisphenoid
Bone of the braincase that forms the floor of the cranial cavity
Single
- Median and mostly ventral/anterior to otic region
- Mostly covered ventrally by the parasphenoid
- Forms basal articulation with the palatal complex via the basipterygoid process
Sphenethmoid
Bone of the braincase that contains the olfactory nerves
Single
- Trough shaped, median ossification
- In sphenoid and ethmoid regions
Basic changes in tetrapod skulls
Early tetrapods
- Skull is formed of many different bones
Later forms
- Many bones are lost or fused, leading to fewer total bones
- Many elements are rearranged
Basal Amniote Skull
Similar to that of an early tetrapod, but with some changes
- Otic notch closed
- Intertemporals lost
- Other temporal bones reduced and displaced posteriorly
- Postparietals and tabular change, eventually become fused to occipital
Pelycosaur Skull
First intermediate stage toward a mammalian skull
- Temporal fenestra is developed
Non-Mammalian Therapsid Skull
Second intermediate stage toward a mammalian skull
- Enlargement of the temporal fenestra
- Quadratojugal reduced and eventually lost
- Squamosal differentiates, with part becoming the zygomatic arch and part helping form the braincase
- Loss of the prefrontals, postfrontals and postorbitals
- Original three skull areas are integrated
Anapsid Skull
- Small brain sits in braincase
- Dermal skull roof covers brain
- No fenestra in skull roof
- Jaw muscles attach to inside of dermal skull roof
Early Synapsid Skull
- Brain somewhat larger than anapsid
- Dermal skull roof covers brain
- Fenestra exists but is covered by heavy connective tissue
- Jaw muscles attach to inside of skull roof and fenestrae
Early Therapsid Skull
- Brain larger than in early synapsids
- Temporal fenestrae have enlarged
- Cranial kinesis is lost