LSM3261 Life Form and Function

Support and locomotion

LSM 3261 Life Form Structure & Function

1st zoology lecture - Animal diversity and basic designs

2nd zoology Lecture - Animal symmetry

Organisation of the animal body; Transmission of messages/materials within the animal body
No. 3 - Protection No. 4 - Support & Locomotion No. 5 - Locomotion (Flight) No. 6 - Sensing the environment, Feeding; Other adaptations
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Animal form and function in relation to:

Objectives

To learn about various concepts and modes of support and locomotion in animals
Hickman et al., 2011. Integrated principles of zoology. 15th Ed. McGraw Hill. Ruppert, Fox & Barnes, 2004. Invertebrate Zoology: A Functional Evolutionary Approach. 7th Ed. Brooks Cole. Pough et al., 2009.Vertebrate Life. 8th Ed. Pearson International Edition. Young, J. Z., 1981. The Life of Vertebrates, 3rd Ed. Oxford. Liem, K. L., W. E. Bemis, W. F. Walker, Jr. and L. Grande, 2001. Functional Anatomy of Vertebrates. An evolutionary Perspective, 3rd Ed. Brooks/Cole.

References (pick out topics in index and contents page):

Locomotion

Major characteristic of animals: nd food, escape predators, nd a mate or habitat. Contraction of muscles results in movement Skeleton supports and transmits movement

I - Support
The role of an antagonist to muscle contraction.

1. Hydrostatic skeleton 2. Exoskeleton (non-living) 3. Endoskeleton (living) 4. Muscular Hydrostats

Support

Bodies of larger animals require support to function normally Soft-bodied aquatic species need to hold themselves up to perform certain functions (e.g. feeding). Dont need to support their body weight Terrestrial species need to support body weight and require stronger support structures
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1. Hydrostatic Skeleton
Found in many soft-bodied animals Fluid held under pressure in body or parts of body Contraction of muscles push xed volume of uid from one part of body to another, transmitting force Hydrostatic skeleton transmits force throughout the animals body changing shape and movement of body
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Hydrostatic skeleton helps in locomotion

Aquatic invertebrates without exoskeleton (e.g. Hydra, jellysh) Aquatic and terrestrial invertebrates with nonliving exoskeleton (e.g. insects, crustaceans, annelids) Aquatic invertebrates with living endoskeleton (e.g. echinoderms)
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1. Hydrostatic Skeleton
a. Hydra

Hydrostatic skeleton? Crude movements only No rened movements

1. Hydrostatic Skeleton
b. Annelids Hydrostatic skeleton?

Body segments separated by septa Fluid compartmentalised in each segment Each segment capable of independent movement Different sets of muscles can act independently in different segments Movement more versatile/rened

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Darwin (1881) burrowing into very compact soils is effected by the worms literally eating their way through it. Newell (1950) Radial, circular, longtitudinal muscles and sphincters around pores. Ideal uids are incompressible, can be distorted totally, and pressure affecting the uid is subsequently exerted in all directions.
View animation at: http:// www.biology.ualberta.c a/courses.hp/zool250/ animations/ Earthworm.swf 11

Earthworms and plant roots both dig through soil

Earthworms and plant root both exert an axial and radial pressures to penetrate soil. The pressures exerted by roots are called growth pressure. By exerting radial pressure, the soil is broken up to allow axial penetration. Radial pressure is always higher than the axial pressure. By exerting a high radial pressure earthworms break up the soil in order to be able to penetrate it with a lower axial pressure.

Axial and radial pressure exerted by earthworms of different ecological groups Keudel, M. & S. Schrader, 1999. Axial and radial pressure exerted by earthworms of different ecological groups. Biology and fertility of soils, 29(3): 262-269.
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Earthworms dig at different depths

Bouch (1977) classied earthworms into ecological groupsanecic, endogeic and epigeic. The endogeic species generated a higher radial pressure because they have a higher burrowing activity due to their geophagous diet.
Keudel, M. & S. Schrader, 1999. Axial and radial pressure exerted by earthworms of different ecological groups. Biol Fertil Soils, 29: 262269.

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Terriswalkeris terraereginae
(Australian endogeic, Family Megascolecidae, 2m)

http://scienceblogs.com/zooillogix/2008/05/ giant_blue_earthworms_and_frie.php

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Giant Gippsland earthworm, Megascolides australis

usually 2-3m, record is 4m

http://scienceblogs.com/zooillogix/2008/05/ giant_blue_earthworms_and_frie.php
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Diplocardia mississippiensis collection in Florida's Apalachicola National Forest

"It is often said that if the ground is beaten or otherwise made to tremble worms will believe that they are pursued by a mole and leave their burrows," Charles Darwin wrote in The Formation of Vegetable Mould through the Action of Worms (1881)
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1. Hydrostatic Skeleton
Hydrostatic mechanism in animals with other types of skeletons c. Echinoderms (e.g., sea stars, sea urchins) - possess calcareous endoskeleton but hydrostatic skeleton moves tube feet

Hydrostatic skeleton?

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Fluid-lled water-vascular system

Ampulla at base of foot

stores uid. contracts, feet

Valve shuts, ampulla uid forced into tube (canals + ampullae +

hydraulically operated, thin-walled tube feet or podia)

Where does the uid come from? What is the madreporite?

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Source: Mechanical Design in Organisms by Stephen A. Wainwright (1982). Princeton Univ., 423pp.

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Hydrostatic skeleton: limitations

Limited to aquatic or soil dwelling animals Water provides buoyancy, soil some support No protective function Not weight bearing: limits size

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2. Non-living Exoskeleton

Secreted by epithelial tissue Supports body, gives body a xed shape Rigid Flexibility in the form of
separate pieces of plates joined by exible joints

Molluscs calcareous shell

What other animals?

Arthropods chitinous cuticle
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Arthropod exoskeleton
What is it made of? What function does it provide the animal with? How is it similar to our bones? Which part of the insect body would you pin an insect through? What is moulting? What is an instar? Are all arthropods groups equally hard? What is the signicance in museum specimens?
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Examples of exoskeletons

Calcareous shells of clams and snails have incomplete exoskeletons for the external shells do not completely cover the animal.! Much movement still requires their hydrostatic skeleton as well so they have both an exoskeleton and a hydrostatic skeleton. Calcareous body coverings of crustaceans (crabs, lobsters, isopods a.k.a. pill bugs).! Crustaceans are arthropods, and like all arthropods, have a jointed exoskeleton. Chitinaeous body coverings of other arthropods (insects, spiders, millipedes, etc.).! When the muscle contracts, the upper part of the exoskeleton is deexed resulting in a downward wind beat.
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Exoskeletons are composed of a range of materials.

Bone, cartilage, or dentine - ostracoderm sh and turtles. Chitin - arthropods, also some fungi and bacteria. Calcium carbonate - shells of molluscs, brachiopods and some tube-building polychaetes. Silica - exoskeleton in microscopic diatoms and radiolaria. Agglutinated exoskeletons by sticking grains of sand and shell to exterior - some formanifera.

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Exoskeleton analogues

Armadillo (constructed from bone) Pangolin (hair) Reptiles

turtle (bone) crocodiles (bony scutes and horny scales)

Echinoderms? What sort of skeleton is the test?

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3. Living Exoskeleton (internal skeleton)

Living (connective tissue) grows with increasing size of

the animal

Different forms:
Almost complete armour just beneath skin (e.g. echinoderms calcium carbonate)

Skeletal framework to support entire body Skeleton built to transmit force (joints) (vertebrates
cartilage/bone)

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Animals with non-living exoskeleton

Armour plates with exible joints used effectively for locomotion

Animals with living endoskeletons Sea urchins, sea stars rigid endoskeleton round body. Hydrostatic skeleton moves tube feet Vertebrates internal skeleton designed for effective movement
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3.1 The Vertebrate Skeleton

Cranium (skull) [sometimes regarded as part of axial skeleton] Visceral skeleton - supporting gills (gill arches), jaws and tongue Axial skeleton (vertebral column, ribs) supporting rest of body Appendicular skeleton (jointed limbs and limb girdles) supporting rest of body

skull

vertebrae

ribs
pectoral fin paired) pelvic fin (paired)
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3.1 The Vertebrate Skeleton

Aquatic species Less need to support body weight Streamlined Terrestrial species (tetrapods) Must support body weight Development of pentadactyl limb
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Aquatic vertebrates

Vertebral column elongated Built to withstand compression Limb girdles reduced

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Terrestrial vertebrates (Tetrapods)

Vertebral column

Shortened Greater rigidity

Appendicular skeleton
Well developed Development of pentadactyl limb

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General shortening of vertebral column as the animal becomes more terrestrial (Part that is involved in locomotion)

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Importance of the rigid vertebral column in tetrapods

The rigid vertebral column in tetrapods, acts as a girder because limbs bear the full body weight (cf. sh: exible, less ossied, constructed to withstand compression during locomotion, while aquatic medium bears body weight) Emphasis on strength and rigidity instead of exibility
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skull

pectoral girdle

pelvic girdle
vertebral column

front limbs
Axial skeleton Skull Vertebral column

hind limbs
Appendicular skeleton Pectoral and pelvic girdles Paired limbs

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Hypothesised stages in evolution of the pentadactyl limb

A skeleton with stronger and more rigid limbs to lift body off the ground (cf. ns) led to the development of pentatdactyl limb. Hypothesised stages in evolution of pentadactyl limb from sh n: (a) Elongation (b) Bending (c) Develop another joint (wrist/ankle) - greater surface area contact with ground for better balance (d) Shifting of limbs from lateral to ventral side for greater leverage (e) Orientation of limbs for speed (lateral view)
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Lightening skeletal weight, particularly in terrestrial animals.

Terrestrial vertebrate (e.g. amphibian):

Flattened with loss/reduction of most dermal bones Large cavities (e.g. orbits). lighter skull No operculum Hind part (in sh, concerned with gills and pharynx) becomes reduced

Cf. Aquatic vertebrate (e.g. sh):

Deep, heavily protected by many dermal bones, large operculum to protect gills. supported by water

Liem et al., 2001

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Similarities between arthropod and vertebrate design - the articulated skeleton

Articulated skeletons - skeletal components meet or articulate at the joints, allowing one part of the body to move in relation to another. Muscles spanning joints and anchored to different parts of the skeleton provide the power for movement.

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Similarities between arthropod and vertebrate design - the articulated skeleton

Articulated skeletons serve two functions.

They allow the retention of a characteristic physical form. They support an organisms weight and resist the stresses of locomotion.
What kind of animal structure provides protection, but does not dene an animals form or support its weight?

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The Articulated Skeleton

Two novel parallels in arthropod and vertebrate evolution -

both are the most successful of all terrestrial animals both are the only organisms ever to evolve an articulated skeleton

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4. Muscular Hydrostats
Densely packed three-dimensional array of muscle bers. Lack the rigid elements of skeletal support systems Lack the uid-lled cavities of hydrostatic skeletons This musculature generates forces for movement, deformation and changes in stiffness and skeletal support.

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4. Muscular Hydrostats
Examples: arms and tentacles of cephalopods, tongues of mammals and lizards, trunk of the elephant

Kier Lab, UNC Chapel Hill

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4. Muscular Hydrostats
Also in manatee, possibly in the dugong also?

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4. Muscular Hydrostats
A dynamic skeletal support system. Rigid skeletal elements are restricted by joints; Muscular hydrostats can deform - bend, elongate, shorten and twist at any location and at multiple locations simultaneously Muscular hydrostats can highly localise deformations, unlike hydrostatic skeletons. E.g. octopus arm or tongue,

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II - Locomotion in water, land and air

Dierent strategies for dierent densities

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Locomotion in water, land and air

Water
No need to support body weight Can remain sessile or sedentary; nekton must be active Need to cope with pressure

Land
Need to support body weight

Air
Need to support body weight Must actively generate propulsive force

Can remain suspended in water column Need to maintain balance in 3D perspective

Need to move in search of food, resources Need to maintain balance (2D medium)

Need to keep moving at all times Must maintain good sense of balance in 3D perspective

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Locomotion in water, land and air

Aquatic medium
! ! ! !

Can remain sessile or sedentary (need holdfast mechanism) Can be strong swimmers Generally carried about by currents Support limbs reduced

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Modes of Swimming
Beating antenna or agella - smaller forms Flex cup shaped bodies or drift (passive) (jellysh etc) Undulation of body (vermiform organisms in many
phyla), arms (feather stars) pleopods or pereiopods)

Paddling with special swimming legs (crustacean Flapping n-like structures, e.g. clapping two shells Propulsion - jet (cephalopod) and tail (sh) Energy demands - buoyancy and sudden movement
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Swimmers

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Stabilisation of sh in a 3D uid medium

Unpaired ns (dorsal and anal ns) counter yaw, roll Paired ns (pectoral and pelvic ns) counter pitch Pectoral ns also used for braking, steering cf. ight in birds later (cf. stiff, inexible pectoral ns of sharks used as hydrofoils)
Side-to-side oscillation of vertebral column
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Terrestrial Medium

Modes of locomotion

Legged sprawling, e.g. lizards, insects, also mudskipper,

octopus

semi-erect, i. e. elevated sprawling, e.g. monitor

lizards, crocodilians

Also limbless, rolling

fully erect, e.g. mammals , birds

Wired 2010

sprawling

semi-sprawled "high walk"

erect

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Belly clear off ground!

Terrestrial animals with skeletons use their limbs to walk,

trot, run, leap, brachiate, and y.

Since arthropods and vertebrates are typically with bellies

clear off the ground, the body passes through air instead of over the ground with friction. The foot of a running arthropod or vertebrate touches the ground long enough to propel the animal forward with each stride. I.e. moves swiftly, cf. ground-bound, soft-bodied animals.
A Vertebrate Looks at Arthropods, by Barbara Terkanian. Arizona-Sonora Desert Museum, c. 2006.

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Crocodiles high walk and belly crawl over land

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Crocodilian locomotion

Floating - hind limbs splayed out in water with toes and toe webbing extended (submerge
backwards by upward movement of spread limbs)

Swimming - limbs held against body, undulation Diving - front limbs lifted almost vertically, protrude above the shoulder to direct head downward
IUCN Crocodile Specialist Group
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Crocodilian locomotion

High walking - most common gait: limbs held erect beneath body, tail drags Sprawling - rapid movement on land: front and back legs on one side meet when body curves, then separate, tail thrashes from side to side in synchrony Sliding - going down steep mud banks: drag limbs, tail moves side to side for propulsion
IUCN Crocodile Specialist Group
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Crocodilian locomotion

Galloping - Australian Freshwater Crocodiles (Crocodylus johnsoni) gallop almost every time they need to move rapidly on land.

Front limbs go out and forward as the hind limbs propel the body forward. Tail tends to move up and down rather than from side to side. Maximum speed about 18 km/h (cf. 2-4 km/h high walk); but exhausted by 100 m

IUCN Crocodile Specialist Group

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III. Adaptations of the pentadactyl limb & vertebrate skeleton

Mammals demonstrate a wide variety of adaptations of the pentadactyl limb and skeleton for dierent modes of locomotion

3.1 Cursorial locomotion in hoofed animals 3.2. Arboreal and terrestrial habits in primates 3.3. Graviportal locomotion in large, heavy terrestrial mammals 3.4. Swimming in aquatic mammals 3.5 Adaptations of vertebral column
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3.1 Cursorial locomotion in ungulates (hoofed animals)

Ability to run fast - another form of defence Skeletal modications of pentadactyl limb:

Limbs lengthened and modied for cursorial locomotion (fast running)

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3.1 Cursorial locomotion in ungulates (hoofed animals)

Skeletal modications of pentadactyl limb:

Elongation of lower

limb segments; shortening of upper segments.

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Skeletal modications of pentadactyl limb:

Locomotion digitigrade (on all digits), developing into

unguligrade (on main digits, others reduced or lost)
Fast running: Distal sections elongated.

http://www.cartage.org.lb/en/themes/sciences/Zoology/ Animalclassication/OrderPrimates/posture.jpg
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3.1 Cursorial locomotion in ungulates (hoofed animals)

Hoof formation - terminal phalanges touching the ground becomes broad, with claw surrounding it The long metapodials (metacarpals and metatarsals) fused together to form a cannon bone (in horses): 5 toes to 3 toes to 1 toe - toenail

Upper limb muscles play an important role in moving the limbs

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Foot = toes to ankles Metatars als

Ankle bones

Plantigrade entire hand/foot in contact with ground. Weight borne by metacarpals (palms) and metatarsals (sole of foot) .

Digitigrade Only digits in contact with ground. Weight borne by digits (ngers and toes).

Unguligrade Tips of main digits in contact with ground. Digitigrade but lateral digits failing to reach ground are reduced or lost.
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Cannon bone

Ungulates
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Cannon bones

eventoed

oddtoed

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Ungulate = a mammal with hooves

Perissodactyla (odd-toed mammals) - horse, tapir, rhinoceros spp. Artiodactyla (even-toed mammals) - all other ungulates.

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The Artiodactyls are more successful than the Perissodactyls

Perissodactyls appeared in late Palaeocene epoch and were most diverse during the Eocene epoch. Artiodactyls appeared in early Eocene and radiated in the Miocene epoch during which the perissodactyls declined. From Cenozoic era to present, Artiodactyla - 36 down to 10 families (81 genera); however, Perissodactyla - 14 down to 3 families (5 genera).
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3.2 Arboreal and terrestrial habits in primates

Primates Apes - great apes - lesser apes Monkeys - old world - new world

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Gibbons, lesser apes

Family Hylobatidae 15 species in four genera: Hylobates, Hoolock, Nomascus and Symphalangus Master brachiators

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3.2 Arboreal and terrestrial habits in primates Modification of pentadactyl limb design in primates arboreal adaptations Pentadactyl plan retained without loss or fusion of bones Hand and foot modied for grasping: opposable thumb (pollex) and toe (hallux) Claws of digits transformed into at nails

Young, 1981
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Tarsier (a Southeast Asian primate)

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Arboreal and terrestrial habits in primates

New World monkeys VS Old World monkeys more arboreal prehensile tails

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Arboreal and terrestrial habits in primates

Large apes not always possible to walk on branches like the monkeys (quadrupedal) Brachiate Instead they swing by their arms which have become longer than their legs Knuckle/st-walking When on the ground, apes cannot remain upright for long need to prop themselves up with their hands in a semierect posture
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Arboreal and terrestrial habits in primates

Gibbons and organutans more arboreal, chimpanzees and gorillas more terrestrial. Feet of chimpanzees and gorillas more adapted for walking broader soles and shorter toes In gorillas, like man, the hallux occupies a position parallel to the other toes

Young, 1981
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Arboreal and terrestrial habits in primates

Adapted for arboreal life

Adapted for terrestrial life

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3.3 Graviportal locomotion in heavy animals

! ! ! !

Large body mass supported mainly by stout pillar-like front legs rear legs push Vertebral column with long neural spines and numerous ribs Heavy head counter-balances body weight, pivoting on front legs Relatively slow movement
Neural spines

Graviportal system
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Graviportal locomotion in heavy animals

Rhinoceros body is graviportal, like that of elephants:
! ! ! !

Vertebral column with long neural spines above the foreleg, numerous ribs, almost reaching pelvis. Whole column forms single girder balanced on the forelegs, The large, heavy head provides a counter-balance to the body weight

Florida Department of Education, Ofce of Educational Technology.

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Graviportal locomotion in heavy animals

! ! !

Stout pillar-like front legs that take the main body weight Hind limbs act to push the animal forwards Tall neural spines and large ribs for attachment of muscles to support front legs Enables heavy animals to climb up hill slopes.

Young, 1981
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Baluchitherium (Indricotherium)
the largest land mammal of all time

http://www.stephanecompoint.com/41,,,3504,en_US.html
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Graviportal locomotion in heavy animals

extinct primitive rhinoceros Indricotherium

extinct perissodactyl

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Graviportal locomotion in heavy animals

" Graviportal locomotion and similar forms evolved independently in unrelated groups of terrestrial animals, resulting in convergence of general form.

Associated with attaining large to huge sizes Living examples: elephant, rhinoceros, giraffe, bison, oxen Was seen in the largest dinosaurs (sauropods).

sauropod
dinosaur

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Graviportal locomotion in heavy animals

Brachiosaurus
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Graviportal locomotion in heavy animals Graviportal locomotion in birds

Graviportal A type of locomotion in which the limbs are large, straight and sturdy and bear a large body mass (e.g. elephant). Usually alludes to slow-moving, weight-bearing terrestrial movement Cf. Cursorial Running locomotion in which the limbs are long and slender, and not associated with bearing a large body mass (e.g. antelope). Alludes to animals adapted to running. Graviportal characters large, heavy leg bones arranged in vertical columns as supports to heavy body mass.
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Graviportal locomotion in heavy animals

Graviportal characters seen in extinct, flightless birds E.g.: elephantbirds (Aepyornithiformes) [Madagascar] moas (Dinornithiformes) [New Zealand] cf. Modern ightless birds (e.g. ostrich), which have cursorial characters (slender legs; reduced toes ! reduced contact with ground while running)

Pough et al., 1990

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3.4 Skeletal adaptations of mammals returning to the aquatic environment Seals, sea lions, walruses, dugongs, manatees, whales, dolphins, porpoises

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Whales, dolphins and porpoises

Flipper-like forelimbs and large dorsal n for stability Tail and dorsal n = neomorphs (new structures evolved in response to returning to water. Skin folds without skeletal support) Only fore ippers have skeletal support ! modied from fore limbs. Hind limbs disappeared completely and pelvic girdle reduced Vertebral column reverts to similar form as shes, i.e. a compression structure for aquatic locomotion. With absence of neck. Movement involves dorso-ventral undulations of vertebral column (cf. shes)

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3.5 Adaptations of vertebral column

Importance of rigid vertebral column

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Extension/exion of vertebral column

Fish-like side-to-side oscillation of vertebral column

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