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Temperature regulation in animals by heat exchange Types of animal responses to change in ambient temperature More accurate terms o Homeotherms Regulators Maintain constant body temperature o Result of internal physiological mechanisms o Poikilotherms Conformers Body temperature fluctuates with environment Can also regulate by changing their environment Most reside in a stable, colder environment More or less constant temperature Based on source of heat o Ectotherm Obtain heat from environment o Endotherms Produce heat internally In cold climate regional heterotherms Reduces metabolic cost Still regulate! Temporal heterotherm Allow temperature to drop hibernation o Heterotherm Produce heat internally Not in all body parts at one! Regional Heterotherms Warm up only parts of their body i.e. warming up of flight muscles During heat loss heat moves from deep parts to skin Body wall thickness, conductance, temperature gradient o Determine speed of loss or gain Physical mechanisms of heat transfer Temperature is the measure of intensity of heat Direction of heat transfer determined by magnitude of temperature o Higher temperature to lower temperature Conduction o Direct physical contact o Rate of transfer proportional to Area of surface contact Temperature gradient Material Q kA(T2 T1 ) l o contact surface area must be normal to direction of heat flow Convection o Transfer of heat by mass flow of fluid, gas or liquid o STILL EMPLOYS CONDUCTION o Very effective means of mass transfer Maintains steep temperature gradient Electromagnetic radiation o No physical contact or carrier fluid to transfer heat o All object emit radiation energy Give heat to others without contact o Intensity increases with Temperature Frequency Against wavelength o In organisms infrared o Q C(T2 T1) Evaporation o ONLY for lowering body temperature in hot environment o Evaporation of water needs a lot of energy Thus, it cools a lot! o Sweat, panting, drooling on the body surface Heat exchange between organism and the environment Total heat = heat produced metabolically + heat gained or lost o Depends on mass and overall specific heat or tissues An organism can gain or lose heat depending on gradient Heat exchange in cold environments Core body temperature of endothermic animals o Range of 30C to 42C birds are higher Thermal Neutral Zone (TNZ) o Specific range in which temperature is kept constant No change in metabolic rate o Lower critical temperature (LCT) Below it maintained through metabolic regulation o Upper critical temperature (UCT) Over it maintained by active heat dissipation Thermal conductance o Heat produced in core transported to surface By convection and conduction o Heat escapes by conduction to environment Air temperature and convection and radiation affect it o H C(TB TA ) b C* aMb mass specific conductance o conductance per unit area decreases with mass exponent -17 Conductance and tolerance to cold o Specific conductance decreases with body size faster than specific heat production Larger animals retain more heat than smaller animals Harder to dissipate heat in warmer climates Control of heat exchange in cold environments o Physiological mechanisms Autonomic control of blood flow to skin Reduced through vasoconstriction cold Increased through vasodilation hot Countercurrent principle Hot arteries surrounded by cold veins Smallest endotherms Higher cardiac output increased frequency Piloerector muscles Blubber changing blood flow through it o Behavioral mechanisms o Choice of habitat Control portion of body surface exposed to environment Heat exchange in warm environments Losing heat against a temperature gradient Evaporative cooling o Skin surface, lung surface, drooling o Body water is lost Thick fur insulates against entry of external heat Deep burrows Internal production of heat Metabolic reactions produce heat as a byproduct o May be insufficient to maintain body temperature Shivering thermogenesis o Contraction of antagonistic muscles Little net movement o All energy released as heat Non-shivering thermogenesis o Brown fat oxidized by respiration uncouple from ATP synth. Cells loaded with fat and mitochondria High supply of blood vessels o All energy dissipated as heat If it can’t keep up Hypothermia Ectotherms do not regulate with metabolic rate no TNZ Thermoregulatory processes Set point temperature slight deviations are immediately sensed Very precise switching on and off of mechanisms o Slight deviations adjustments of conductance o Outside TNZ increase in metabolic rate o Extreme thermogenesis pr evaporative cooling Hypothalamus temperature control sensor o Group stimulates thermogenesis, vasoconstriction,piloerection o Group stimulates vasodilation, sweating o Group decreases thermogenesis, vasoconstrition, piloerection Biomechanics: structural support and organismal body Principles of mechanics Forces acting on a body o 3 major forces Compressive force Tensile force Shear force o Stress is caused by these forces Deformation it causes strain Depends on magnitude of force & material o Strength Capacity of the body to resist force without breaking or buckling of permanent damage Can be measured in compression or tension Young’s modulus of Elasticity o Ratio between stress and strain constant o E quantifies the stiffness of the material Rubber = lowest, Diamond = highest Bone is in the middle Elastic and plastic strain o Strain is only proportional to stress in true hard objects o Cartilage does not obey Hooke’s law o Elastic strain completely reversible Linear with stress in region of elastic response o Plastic strain permanent and non-reversible After the yield point, then fractures Cantilevers only supported at one end o Strut is under compressive o Stay is over tensile No stress in the middle part beam can be hollow Skeletons More or less firm structures that maintain structural integrity Provide attachment sites for muscles Hydrostactic skeletons o Fluid-filled body cavity surrounded by circular and longitudinal muscles o Earthworm o Plants also have a hydrostatic skeleton Exoskeleton o Generated by secretion to outer surface hardens o Muscles attached to inside o Gives shape, anchorage, protects 1.135 o Mskel 0.078Mb o M shell 0.0482M egg 1.132 F 50.86Mb 0.915 Endoskeletons o Internal framework of bones o Axial and appendicular skeleton Composition of skeleton 2 types of connective tissue o Dispersed cells in a extracellular matrix o Cartilage Chondrocytes secrete collagen, elastin, polysaccharides Collagen reinforce gel matrix Flexible & resistant to compression Lines the joints & flexible external support o Bone Osteoblasts Secrete extracellular matrix collagen + Calcium When trapped osteocytes Osteoclast resorbs matrix (erodes bone) Bone is constantly replaced and remodeled Perceive stress imposed on them & strengthen structure Calcium is also a reservoir for the body Marrow cavity in the center Cancellous ends spongy looking Compact bone Haversian systems (osteon) Series of concentric layers with canal in center Occupied by blood vessels and nerve Canals parallel to long axis Interconnected by Wolkmann’s canals Orientation of collagen alternate between lamellae Osteocytes present in central canal The skeleton as a supporting system Strength and design of bones o Can take more compressive than tensile stress o Ends are wider than the main shaft o Forms joints with other bones Lined with cartilage for smoothness, lubrication, stiffness and resiliency o Spongy material trabeculae Struts going from one edge to the other o Thin bones compact on outside & cancellous on inside Allometry of bones o Bones of large animals scaled out of proportion to dimensions o Area of bones has to increase in proportion to weight o Large vertebrates must be aquatic buoyancy of water o Skeleton has to support static weight and locomotion Bones as beams and columns o Only true for graviportal animals very large and heavy More or less vertical columns solid without marrow Made for supporting not running o In eursorial animals successive limb bones at an angle Subjected to multidirectional stress Hollow or marrow cavity Distal end compressive stress Proximal end tensile stress Skeleton as a balanced structural frame Keeps the animal from collapsing and protects against toppling Cantilevered mass is balanced out by the other side Neck and head cantilever supported by nuchal ligament o Most elastic part of the body Allometry of trees Base diameter has to increase to 1.5 power of height Heights of all trees are below critical buckling line sense stresses Terrestrial Locomotion Muscle and motion 3 types of muscles o Cardiac muscle Looks striated Arrangement of actin and myosin filaments Criss-crossing network Protects heart from tearing Can withstand high pressure Electrical contact with each other Pacemaker cells o Smooth muscle Not striated Ling and spindle-shaped with one nucleus Force of contraction of internal organs Arterioles & veinules wrapped by single muscles cells Nerve cells between muscle layers Direct control by nervous system o Skeletal muscles Involved in all voluntary movement Striated and have several nuclei Formed by fusion of cells Much larger than 2 other types Several bundles of muscle fibers Many myofibrils Bundle of thin actin and thick myosin Divided into contractile units sarcomeres o Cylindrical with Z-line at both ends o Contraction sarcomere shortens o Mimimum contraction twitch Contraction by firing of motor neurons Depends on number firing and frequency of firing Spatial and temporal summation Slow-twitch fibers red muscle Lots of myoglobin, mitochondria and blood vessels High level of ATP, glycogen and fat Long term aerobic work Maximum tension is lower Fast-twitch fibers white muscle Less myoglobin, mitochondria and blood vessels Develop higher tension and quicker Cannot sustain it for long sprinting Powered by glycolysis (anaerobic respiration) Muscle-bone attachment o Attached to bone by tendons o Bone to bone by ligaments Principle of lever and bone-muscle machines Principle of levers o 3 point of a lever Fulcrum pivoting point Effort arm Load arm o Class-1 lever Fulcrum between effort and load arm See-saw o Class-2 lever Load arm between fulcrum and effort arm Wheelbarrow o Class-3 lever Effort arm between fulcrum and load arm o Effort x Effort Arm = Load x Load Arm o Lever rotates in direction of greater torque o Effort needed inversely proportional to length of effort arm o Effort arm short compared to load arm quicker! effort velocity x load arm = load velocity x effort arm Structural adaptations for fast running Cursorial animals adapted for high-speed running Non-cursorial animals adapted for walking not running Speed of running determined by stride length & frequency Limb length and foot posture o Speed can be increased by increasing limb length Elongation of foot bones o Plantigrade whole foot o Digitigrade ball of foot o Unguligrade toe tips Shoulder position o Non-cursorial scapulas horizontal with clavicle o Cursorial vertical scapulas without clavicle Rotation in same plane and direction as limb More freedom of movement without clavicle Carnivore have a reduced clavicle Movement of the spine o Legs are under the body spine is free to flex in vertical plane o Stride length is increased with right flexion and extension Allows extra rotation of hip and shoulder girdles Legs reach out farther, front and back o Bounding also adds to stride length Angle and speed of motion o Greater swing of the limb when muscle attached close to joint o Higher load arm to effort arm ratio more speed, less power Limb mass o Limbs move forward and backwards during locomotion Inertia affects them Proximal distribution of limb mass decreases inertia o All muscles involved in running located near the torso o Some bones at distal end reduced or eliminated completely Special locomotion modes in hot desert o As small a body part as possible touching hot surface for as little time as possible Tiptoe stance in lizards Sidewinding snakes Cartwheeling spider Swimming and Flying Principles of fluid dynamics Forces against forward motion o Drag o Cross-sectional area perpendicular to flow resistance Laminar and turbulent flow o Laminar flow smooth flow pattern Parallel to direction of flow Narrow and short wake o Turbulent flow rough and disorganized flow pattern Perpendicular to direction of flow o o o o Swirling eddies behind plate Pressure drag Against the forward movement Depends on cross-section, density and velocity Fiction drag Depends on surface properties of plate & viscosity Streamlining Tapering of the trailing surface of a sphere Fitness ratio Ratio of maximum thickness to chord length Turbulence results when flow separates from surface of body Reynolds Number (Re) length speed density o Re dimensionless number density Boundary layer o Velocity flow is zero at interface between fluid and surface o Boundary layer thickness inversely proportional to flow speed Lift to Drag Ratio, Angle of Attack and Aspect Ratio o Fluid moving past bottom positive pressure +upward thrust o Fluid moving past upper surface low pressure o Center line or camber line Equidistant from upper and lower of the airfoil o Chord line Connects centre of leading edge to that of trailing edge o Camber o o o o High o o Distance between center line and chord line Magnitude of lift depends on angle of attack Maximum lift at 10-20 degrees lift : drag ratio can’t be increased only by surface area arrangement! Length : width ratio aspect ratio Streamlined body has less drag than rectangular body lift and anti-stalling devices Lift force can be increased increasing camber, thickness, SA Lift increases then decreases with angle of attack Stall decrease of air flow slow over airfoil Flow separates from surface Coefficient of drag exponentially increases with a of a Coefficient of lift increases then decreases and gaps as anti-stalling devices Redirects air flow from lower to upper surface o Slots o Bird Have alula which pops up just before landing Also have slots Stability and control yaw, pitch and roll o Yaw Side turning around vertical axis Corrected by vertical rudder o Pitch Up and down rotation around horizontal axis Corrected by horizontal stabilizer o Roll Turning movement around longitudinal axis Corrected by ailerons Speed and drag Re determines relation between drag, speed and body size o Low Re friction drag predominant o High Re pressure drag predominant Swimming Size, speed and metabolic cost o Fish have same density as water naturally buoyant Only have pressure and friction drag as resistance o Df 0.5SU 2C f Drag increases with square of velocity o U 19.5(length) 0.5 M 0.17 larger fish swim faster than smaller fish MSMC decreases with increase in body mass Propulsion mechanisms in fish swimming: Undulatory swimming o Zigzag lines separate muscle blocks myotomes Segments along length of fish with vertebrae Cause a wave of undulation along sides of body Start at anterior end and passes down to tail o Tail propulsion Shows largest displacement specialized organ Adaptations for maintaining buoyancy and reducing drag o Swim bladders Beneath vertebral column Expels and absorbs gasses regulates volume Always match density of surroundings o Fish slime and drag Secrete mucus on outer surface of skin Protects against parasites and reduces drag! o Tuna as super-streamlined body Smoothness of skin no mucus Fins are thin and can be tucked into a groove Eyes flush with surface Narrow caudal fin is good for cruising Flying Flying ability and body size o Wing-loading is proportional to length o Stalling speed proportional to square-root of wing-loading Larger birds higher stalling and take off speed Running start, wind, higher launch, etc Effortless flight thermal air masses Small fliers (bees) rapid wing beat + vertical take off Some can hover at will o Wing characteristics Aspect ratio Length and width of wing High aspect ratio = high lift to drag ratio Can glide at more acute angles close to ground Flapping of wings Downward and forward power stroke Counteracts gravity and drag Backward recovery stroke Wing partially retracted In hummingbird lift also generated Wings rotate during recovery Wing-loading Ratio between weight and wing SA Lower wing loading o Lower stalling speed o Enables tighter turns Soaring and gliding o Soaring Altitude is gained or conserved thanks to thermals Metabolic cost adjustment of tail & wing position o Gliding Altitude is lost, however slowly Hormones in animals Cellular secretions Hormones are chemical messengers and regulators o Produced at on location and act on another o Effective at very low concentrations Autocrine secretions o Act on the cells which produces them o Secreted outside and bind to receptor on plasma membrane Paracrine secretions o Secretions from one cells affect cells surrounding Endocrine secretions o Poured into and transported by blood stream o Act on cells of distant organs and tissues o Clustered together to form gland Exocrine secretions o Release outside the body of the organism Skin, epithelial surface of the gut, etc o Secreted from a gland Duct that carries the secretion Neurosecretory cells o When secreting cells are hormones o Almost all hormones o All neurosecretory hormones are proteins Neuropeptides Types of hormones Peptide hormones o 3-4 AA residues to very large proteins o water soluble cannot diffuse into cell o specific membrane-anchored receptors o initiate cascade of signaling events Steroid hormones o synthesized from cholesterol o lipid soluble can diffuse into cell o receptors in cytoplasm o sex hormones testosterone & progesterone o Receptor-hormone stimulates or inhibits gene expression Catecholamine hormones o Derived from modification of amino acids o Stress response epinephrine and norepinephrine o Hormone binding causes change in membrane potential or triggers second-messenger pathway Eicosanoid hormones o Prostaglandins Lipid-soluble and lipid-insoluble hormones o Signaling mechanism depends on solubility and partitioning SEE PAGE 92 OF COURSE PACK!!! AND MEMORIZE Hypothalamus and Pituitary gland – A major control center Hypothalamus is lower region of the brain pituitary right beneath o Anterior and posterior pituitary Neurosecretory cells of hypothalamus axons into posterior o Main cell bodies in the hypothalamus o Synthesize hormones Control synthesis and secretions of hormonespituitary Tropic hormones Hormones released by hypothalamus o Stimulatory and inhibitory effect on anterior pituitary gland o TRH, GnRH, Prolactin-IH, ProlactinRH, etc… P.94 Hormones released by pituitary gland o Many are tropic in nature o Main function of posterior Secrete 2 hormones Oxytocin labor in mammals ADH water balance osmoreceptors in hypothalamus pressure receptors in aorta Vertebrate endocrine glands and other hormone secretors Adrenal Glands o Adrenal medulla inner part Develops from nervous tissue Controlled by nervous system Secretes epinephrine and norepinephrine Epinephrine also acts as neurotransmitter o Adrenal cortex outer part Secretes aldosterone, cortisol and cortisterone Play roles salt and carbohydrate metabolism Controlled by tropic hormones Heart atrium o Secretes atril natriuretic peptide (ANP) Controls salt and water excretion by kidney Kidney o Secrete hormone calcitrol Regulates Ca level in blood, bone formation and absorption of calcium and phosphate in intestine Pancreas o Alpha cells secrete glucagon o Beta cells secrete insulin o Regulate blood level of glucose in opposite manner Parathyroid gland o Secretes hormone parathormone Regulates blood levels of calcium and phosphate Pineal or epiphysis o Secretes melatonin Role in photoperiodic effects Anti-gonadotropic action Thymus gland o Secretes thymic hormones Regulate proliferation and differentiation of lymphocytes Thyroid gland o Follicular cells secrete 2 hormones Thyroxine and tri-iodothyronine Regulation of growth and differentiation, metabolic rate and consumption of O2 and heat production o Parafollicular cells clusters called ultimobrachial glands Secrete calcitronin Down regulates calcium levels in blood Hormones involved in digestion SEE PAGE 99 Pheromones, allomones and kairomones Pheromones o Used to communicate with members of same species o Pheromones evolved into hormones o Exocrine secretions o Involved in mate location, alarm and alerting, defense behavior and establishment of caste system o First pheromone identified bombykol o Can respond to a single molecule o Quite unique molecules o Odor-plume is produced and travel down wind several km Active space o Can be used as odor-trails Allomones o Adaptively favourable to the animal that produces them o One-way chemical weapons which attack receiver o 2-way where both the receiver and emitter benefit Kairomones o Favourable to receiver but harmful effects on emitter If becomes attractant to predators or parasites Evolutionary perspectives on hormones o Much biochemical unity in different life forms o Similarity in amino acid sequence o Hormones in plants Special features of plant hormones Major differences with animals on sources and actions of hormones Nothing comes from a gland Some are synthesized in particular part of body Specific action is not well defined for most o Interact with each other to produce spectrum of effects o Some are more specific than others Concentration in cell determined by o Rate of synthesis o Rate of degradation o Rate of sequestration Binding to other cell molecules or compartmentalization Auxins 3 types of auxins exist in plants Stimulation of growth by elongation Formation of adventitious and side roots Maintenance of apical dominance o Growth inhibition of lateral branches from main axis of stem Inhibition of leaf abscission Coleoptile tip exposed to light makes shoot bend toward light o Auxin mediates phototropism o Auxin is transported to shaded side Cells grow faster and cause bending towards light Transported basipetally o From morphologically upper to morphologically lower Polar transport mediated by auxin efflux carriers o Anchored in PM only on morphological lower side of each cell Gibberellins (GAs) Terpenoid compounds More than 100 gibberellins are known o Only a few have potent biological activity (GA3 is most used) Role in seed germination and stem growth o Dwarf pees have mutation in GA production gene o Applying GA will restore wild type phenotype Cytokinins Stimulation of cell division Lateral-shoot induction Retardation of senescence Abscisic acid (ABA) Synthesized in response to dehydration of plant tissues Seed MUST dehydrate and accumulate ABA o Without ABA vivipary Precocious germination on the mother plant Immediate precursor ABA aldehyde o Inability to produce reduced drought resistance Ethylene Synthesized from methionine Fruit-ripening hormone Autocatalytic o Any ripe fruit will make other fruits ripen faster Can be oxidized to ethylene oxide Brassinolide Only truly steroid hormone in plants Multiple effects on growth and development Systemide Protein hormone Synthesized in response to insect attacks Initiates synthesis of protease inhibitors I and II o Prevent digestion of proteins by insect Salicylic acid and Jasmonic acid Response to fungal, bacterial and nematode pathogens Can become volatile by methylation Growth and development 1 – General considerations Atomic size and organismal size Biological systems have predictable and accurate activities o Very small margin or error governed by sqrt(n) largers numbers = larger accuracy, stability, predictable Measurable number of proteins are defective o Mechanisms to evolve and degrade them and recycle AA Growth Irreversible increases in size or volume or an organism o Unlimited resources N t N oe kt ln Nt kt ln No o Limited resources dN kN(N max N) Nmax-N = growth to be achieved dt S-shaped curve Lag period Log phase Stationary phase Dynamic nature of organisms Organism is constantly changing o Starts as single cell Undergoes morphologically different and increasingly complex stages of development Haploid gametes diploid organism Cellular basis of growth and development (Cell division and enlargement) Multicellular organisms also build body structures Cell division is basis for growth AND development o Provides growth through increasing cell numbers o Provides basis for cell commitment and determination o Complex and highly regulated process o Initiated by signals received by the cell o Karyokinesis division of nucleus o Cytokinesis division of cytoplasm Rest of time interphase (between divisions) o Cells receives & responds to biochemical signals Regulate the progression of cell division process o Cell enlarges before it divides! Development: Cell commitment and cell differentiation Division then differentiation basis of organismal development Continuous supply of cells is needed o Repair and replaced of worn out cells Stem cell (meristematic) o Unspecialized or uncommitted cell o Capable of cell division o Can differentiate into any type of cell Differentiation o Cell commitment, determination & specialization Has specific structure and function o Incapable of cell division o 3 levels Structural Morphology and anatomy Biochemical New enzyme activity or gene products Molecular Gene activation or repression Development o Qualitative changes during the life cycle Molecular genetic basis of development Genetic constitution determines everything in organism Gene products are present in cytoplasm o Long-lived and stable Generation of polarity as a basis for development Used to achieve a specific shape Unequal cell division o Results in smaller and bigger cell o Common in plants Position of nucleus o Common in lower plants Gradient of morphogens o Proteins, mRNAs, other biochemical factors o Bicoid anterior and nanos posterior Sperm point of entry o Makes that part of egg ventral half of embryo Incident light o Fucus shaded side produces rhizoid cell Positional information and intercellular communication Tissues cells belong to same type & have identical features Cells under influence of neighbours Cell signaling! o Diffusible chemicals Released from one & bind to receptors on other Generates second messenger info to nucleus o Direct contact between surface proteins Leads to generation of second messenger o Gap junctions or plasmodesmata Opening between 2 cells cytoplasms in contact Can transfer large molecules proteins Growth regulators and hormones crucial part o Communication and development Reversibility of differentiation, totipotency of cells and regeneration of missing part Cells cannot de-differentiate o If they did No signals to guide it Become a cancerous mass of cells Uncontrolled cell proliferation Growth and development 2 – Plants Plants growth and it’s measurement Higher plants are fixed in soil o 70% of growth is cell expansion & 30% is cell division Growth can be measure as increase in height or dry weight o Growth curve is typical S-shaped curve LogL2 LogL1 o Relative growth rate t 2 t1 o Rate of cell growth (G) E(P Y) Growth regulation by hormones growth in major way 2 hormones regulate o Gibberellin (GA) Only to intact plant Increasing growth with increasing concentration GA3 is rapidly metabolized Cause hydrolysis of starch Increases glucose concentration Lower water potential Water influx! Increases the term P in equation o Auxin (IAA) Stimulates growth of only excised segments Low concentration stimulates, high inhibits Short lag-time before growth is observed Activates proton-pumping ATPase pH in cell walls declines activates certain enzymes expansins hydrolyze acid-labile bonds cell wall more extensible less pressure needed to expand cell Increases E and decreases Y Growth regulation by environmental factors Inhibitory effects of light on growth o Blue and red are inhibitory Red light can be inhibited by far-red light flash or GA Light receptor phytochrome Effect of blue light is more rapid Cannot be countered Light receptor cryptochrome Regulation of plant growth by plant water status o Water deficit is strong inhibitor o Rapid, direct and reversible effect Plant development: Seed dormancy and germination Seed dormancy o NOT DEAD! o Many seeds unable to germinate right after formation Too strong or impervious seed coat Hormonal or chemical inhibitors Leaks out during rain or water intake o Soaks up water but does not germinate Becomes metabolically active o Activated embryo produces GA promotes germination o Germination balanced by ratio of GA:ABA o Can also require environmental conditions light or chilling Light could mean near soil surface photosynthesis Red light promotes germination, far-red counters Last type of light will determine germination Seed germination o Sequence of events Water absorption, increase in energy charge, mobilization of food reserves, protrusions of root, etc. Water absorption o Mature seeds are dehydration 7-15% weight o Membranes are not lipid bilayer conformation Take time to regain normal characteristics Small solutes leak out o Membrane regain semi-permeability Reabsorbtion of solutes o Bi-phasic process Pre-existing cells absorb water (1st phase) Mew cells are formed and absorb water (2nd phase) Increase in energy charge (EC) o Proportion of phosphate bonds in all AMP, ATP and ADP ATP has 2, ADP had 1, AMP has none EC [ATP] 1 [ADP] 2 [ATP] [ADP] [AMP] Production of hydrolytic enzymes and mobilization of food reserves o Enzymes to degrade carbohydrates, lipids and proteins Amylases, lipases and proteases o Genes for these are activated by GA o o o o o Embryo synthesizes GA GA diffuses to aleurone cells & activates gene transcription Enzymes secreted into endosperm Reserves hydrolyzed into monomers diffuse to embryo Serve as respiratory substrates & building blocks Plant development: Meristems (embryonic regions) Permanent meristems o Continuously produce branches, leaves and roots o Located at particular sites Apical Tip of the shoot Leaves and branches Root apex Roots Lateral Cambium & cambium cork Determinate meristems o Leaf primordia o Flower primordia Can also be intercalary at the base of the leaf in grasses Plant development: Organogenesis of shoot (stem, branch, leaf) Shoot and root apical meristems established in embryogenesis o First cell division unequal long basal cell + small terminal Terminal cell is embryo o Leaves have buds in their axilsprimordial of future branches Upper cells are organized, lower cells are random Cells in meristem never differentiate o Products of their division produce lateral organs & internal o Peripheral zone produce lateral organs leaves & branches Outermost epidermis Inner internal tissues Rib zone vasculature Central zone meristem o Apical no hormones Sub-apical hormones Positional information determines what a primordium is going to do Plant development: Organogenesis of root No sub-apical meristem Root cap in meristem o Sheds peripheral cells o Protects root apical meristem o Secretes many biochemicals into soil Quiescent center in meristem o Reserve apical meristem Cells elongation zone behind meristem o Frequency of division declines Maturation zone behind elongation zone o Differentiation of cells Totipotency of plant cells Cells can de-differentiate and grow completely new plants Differentiation is reversible in plants! Hormonal regulation of differentiation into roots and shoots Mass of undifferentiated cells callus Auxin and cytokinins levels regulate differentiation Regeneration of missing parts High cytokinin (from roots) shoots High auxin (from top) roots Growth and development 3 - Animals 4/12/2010 8:50:00 PM Growth of the body Growth spurt at puberty o On sensing of first signal of sexual maturity rush for max Significant changes to relative proportions of different body parts Tension-driven muscle growth o Linear growth of muscle is couple with growth of bone Driving force is the tension bone exerts on muscle Genetic basis of limb growth o Limb contains genetic information about maximum length Cell division and limits to cell proliferation Contact inhibition of cell division Cells have to be anchored to divide basal membrane Cells away from basal layer are differentiated and don’t divide Establishment of polarity Earliest acting morphogens are usually maternally produced Protein gradient results from polar distribution of their mRNA o Embryogenesis proceeds with precise timing Morphogens are all transcription factors o Proteins that bind to DNA and activate transcription Specific genes involved in developmental process Ventral side is marked by sperm entry point Dorsal side is marked by presence of Nieuwkoop centre Cell commitment and determination How do you know? o Take a cell and transplant it in another region If it forms structure it should have differentiated Cell type identity and unique surface features In every case, cells sort themselves out according to their type o Reorganize themselves like in the embryo Different cell types have different surface features o Including specific proteins and glycoproteins Early embryogenesis: establishment of three primary embryonic cell layers Cleavage by cell division starts soon after fertilization At 16-cell stage division without enlargement o Blastula stage o 3 cell types and cavity blastocoel Further cell divisions and movement create gastrula Organogenesis from the three primary embryonic cell layers In last stages of gastrula blastocoel shrinks o Archenteron takes it’s place forms gut Ectoderm o Outside layer o Epidermis, eye lens, brain, secretory glands, spinal cord, etc Endoderm o Inside layer o Bones,cartilage,connective tissue, skeletal muscles, heart, etc Mesoderm o Middle layer o Liver, pancreas, gut, lungs, pharynx, thyroid, pharynx, etc Primordial germ cells are separated from the rest o Will form sperms and eggs o Protects them from cumulative mutations genetic constancy Totipotency and capacity for regeneration of missing parts Differentiated animal cells are totipotent in nuclear transplant o Need a receptive female for artificial embryo o Not exactly totipotency Regeneration of missing parts o Seen in lower animals o Flat worm will regenerate head and tail Morphogen gradient! o Newt regenerate amputated limb Presence of nerve is regulating factor No nerve = no regeneration Unless unnerved in early embryo Major differences between animals and plants Growth habit o Animals determinate o Plants indeterminate Development o Partitioning of germ and somatic cell lines Animals very early in embryo development Plants none until flowering o Role of gametophyte Animals represented by gametes themselves Plants multicellular embryo sac o Post-embryonic development Animals most adult organs formed in embryogenesis Plants from meristems adult organs develop post. o Cell movement during development Animals cells migrate to where they form an organ Plants cells walls cement cells in place of formation o Regeneration and Totipotency Animals cells have finite lifetime Plants de-differention of cells in vitro o Variety of Organs and cell types Animals hundreds of cell types Plants 40 different cell types with little difference o Hormone synthesis/action, specificity Animals specific site of synthesis, action and function Plants synthesized and acts at many location depends on relative concentration Plant reproduction: options and timing Asexual options for reproduction Through any vegetative part that can give rise to a new plant Does not involve chromosomal rearrangement o Leads to inbreeding depression of fitness Readiness to flower Plants will not flower in juvenile phase Passage to reproductive phase comes after a certain time o Can be marked by changes in leaf shape or other things Plant dies after flowering and seeding Transition of shoot apex from vegetative state to reproductive state Important changes in shape and activity of apical meristem o During transition from vegetative to reproductive phase Increase in diameter or length or both in shoot apical meristem Flower structure and development Almost all flowers have 4 flower organs o Sepals, petal, stamens and carpels Stamen has filament and anther on top Carpel ovary at bottom, stigma on top, style in middle More than one fusion into multi-chambered Flower development o Only A sepals o A and B petal o B and C stamens o Only C carpel o A & C complementary mutually exclusive If C is mutated, A will take it’s place and vice versa Regulation of flowering time – Some general comments Environmental cues involved in regulating time of flowering o Not temperature too variable in day o Relative length of day and night! How to measure flowering response? o Time taken for 1st flower to appear o Number of nodes that bear flowers or inflorescnces o Number of flowers per plant o Percentage of plants that flower in a population Photoperiodic regulation of flowering time Short day plants (SDP) o Flowers when day length is shorter than critical photoperiod o Does not flower when above CP Long day plants (LDP) o Flowers when day length is longer than critical photoperiod o Does not flower below CP Day-neutral plants (DNP) o Don’t respond to day length o Flower after producing a certain number of leaves Related to age of the plant Inductive photoperiod o Combination of day and night length that induces flowering Qualitative and quantitative photoperiodic responses o Qualitative not flower without inductive photoperiod Most SDP o Quantitative flowers slower and less without IP Most LDP Number of induction cycles depends on plant (1 to several) Once flowering is induced cannot be stopped! It is actually night that is critical for flowering response o SDP are long night plants (LNP) o LDP are short night plants (SNP) Non-inductive cycle cancers an inductive cycle Hormonal regulation of flowering time Effective for inducing flowering is GA o Can make LDP flower is SD conditions Temperature regulation of flowering time – Vernalization Some plants need a chilling treatment to flower o Gene (suppresses flowering) is switched off permanently A flowering hormone (florigen)? Perception of photoperiod occurs in leaf but flowering shoot apex o Even if single leaf exposed to IP flowering Hypothetical hormone Florigen o Can be transported through graft union of 2 plants Underlying mechanisms LDP in inductive conditions o Gene called Flowering locus T induced in leaf phloem Protein coded by it transported to shoot tip by phloem Complexes with another protein to initiate process SDP o Hd3a is kept repressed by another gene o In inductive conditions repressor gene is inactivated Hd3a free to express itself Acts like FT protein Pollination and seed development Sex systems in plants Distribution of sexes at level of flower, plant or population Flower o Bisexual, female or male Plant o Only bisexual flowers hermaphrodite o Male and female flowers on same plants monoecious o Male and female flowers on separate plants dioecious Population o Any combination of types of plants Gametogenesis Gametes are haploid Stamen diploid Anther microspore mother cell diploid microspores haploid Ovary megaspore mother cell diploid megaspores haploid Development of pollen grains o Anther has 4 chambers Lining of chambers --. Tapetum payer Cells lyse to provide nutrition to microspore o Microspores mature into pollen grains Becomes multicellular male gametophyte Vegetative and generative cell Shed and dispersed Development of the egg o Develops in the ovule o Mother cell is differentiated Produces 4 megaspores 3 die Nucleus forms 8 nuclei 3 mitotic divisions 4 on each end 1 from each side fuse in center secondary nucleus Middle of 3 at microphyler end egg Other 2 are synergids 3 others antipodals pollen tube enters through microphyle Pollination: Self- and cross-pollination o Various vectors of dispersion Wind Inconspicuous or absent petals Large anthers and stigma Animals Large, showy, fragrant Bees most efficient pollinators Blue or yellow Lines or markings as honey guides UV light flower markings Beetles Strong yeasty, spicy or fruity odor No nectar food storing cells in petals Moths Yellow or white flowers stand out at night Birds Long tubular flower structure for beak Bats Forage at night o Nectar compositions Sugars, org. acids, volatile oils, proteins, enzymes, etc o Sugar concentration Depends on flower and time of day o Special case the fig and the wasp Male synconium Male flowers with mature pollen Just inside the ostiole Female synconium Female flowers line inner wall of fruit Long-styled carpels and short-styled carpels Female wasp with fertilized eggs and pollen enters female synconium lays eggs and pollinates flower Eggs in short-styled ovaries Larvae hatch and eat short-styled flowers Male and female wasps mate Females leave synconium and carry pollen away MALES NEVER SEE LIGHT OF DAY o Advantages of cross-pollination Promotes heterozygosity and genetic diversity Self-pollination Needs self-compatibility and temporal coincidence Promotes homozygosity + inbreeding depression Germination of pollen grains o Pollination pollen grain lands on stigma If compatible absorb water and swells Germinates and produces pollen tube Incompatible no germination If germinatesgrow short distance and dies Sphorophytic incompatibility o Ligand on surface of pollen binds receptor kinases Complete inhibition of germination Gametophytic self-incompatibility o Controlled by several alleles of S-gene o S-RNAse produced by cells lining pathway of pollen tube Pollen tube from self-pollen takes S-RNAse Degrades the mRNAs in pollen tube dies Double fertilization o Compatible pollen hydrates & germinates pollen tube o Generative cell divides mitotically 2 sperms Released into embryo sac One fuses with egg, other with secondary nucleus o Resulting triploid cell forms endosperm o Sinergids and antipodals degrade realease nutrients Development of embryo o 1st division is unequal Smaller embryo Larger suspensor Seed development o Different parts of ovule and embryo sac give rise to different parts of seed Precocious germination o ABA prevents germination inside the mother plant o 2 mutations can lead to precocious germination Inability to synthesize ABA Germinates right in fruit Insensitivity to ABA Mutation in receptor that senses ABA o Viviparous mutants Sequence of events in seed development o Embryogenesis o Seed development, growth, germination genes not repressed Germination is inhibited by ABA o Seed starts drying out DNA synthesis stops o Seed dries out dormant cell Storage food reserves o Starch, proteins, fats, phytic acid or phytate Storage of minerals o Phytate Hexaphosphoric acid ester of myo-inositol phytin Insoluble mixed salts K, Mg, Ca o Storage proteins Albumins Globulins Glutelins Prolamins Reproduction in animals Reproduction and genetic recombination Reproduction increase of number of individuals o By production of offspring Genetics determine potential, environment determines actual In evolution o Reproduction without recombination o Then reproduction and recombination separated in time o Then melding of the two Differentiation of sexes ensures re-shuffling of gene combinations o Enlarges repertoire or responses to given situation Asexual reproduction Single cell and some multicellular splitting in 2 binary fission o Single celled genetic material divided in 2 o Multicellular no chromosomal duplication Genetic constancy is not disturbed Budding small outgrowths produced by main body detach o Start independent existence Chances for genetic innovation are slim Resort to genetic recombination under stressful conditions Recombination through mating without reproduction Macronucleus and all but one micronucleus disintegrate o Undergoes meiosis 4 haploid micronuclei3 die, 1 survives Remaining mitosis 2 micronuclei Exchange of 1 micronuclei between Paramecium Separation of organisms & fusion of micronuclei NO increase in number of cells Asexual reproduction is good under constant environment Parthogenesis o Sexual behavior exhibited but no egg fertilization Unfertilized egg is stimulated to form embryo o Whiptail lizard Acts like female in high estrogen & male progesterone Behavior of courtship display but no sperm produced Essential for egg development Asexual reproduction important in parasites o Tapeworm oldest segment breaks away and becomes adult Sexual reproduction Combines genetic recombination with reproduction o Contributes to generation of genetic diversity o Formation of haploid gametescrossing-over in chromosomes o Assortment of chromosomes is also independent Hermaphroditism o Organism is both female and male both sex organs Origin of internal sex organs o Common origin and represent 2 extremes of hormonal ratios o Structural deviations are due to hormonal balances o 3 major steps Gametogenesis haploid cells produced Mating gametes are brought together Fertilization gametes fuse to form zygote Gametogenesis o In early stages of embryogenesis germ cells set apart Migrate to gonads when later are differentiated Protects germ cells from mutations Reduces birth defects Males sperm, female egg Proliferate mitosis oogonia (female) & spermatogonia Spermatogenesis spermatocytes undergo meiosis Oogenesis oocyte undergoes unequal meiosis Very small cell is 1st polar body discarded Second meiosis second polar body produced o o o o Large cell forms egg Structure of ovary and egg o Ovary contains primary oocytes form definite # of eggs Most degenerate o Each is surrounded by layer of ovarian cells Follicle o Ovarian cycle cycle of ovulation, duration varies with specie Several follicles mature only one releases ovum Follicle cells form corpus luteum Produces and secretes progesterone & estrogen o Egg is surrounded by plasma membrane Just inside PM, cortical granules Egg membrane surrounded by vitelline envelope In turn surrounded by coat of jelly Sperm has to go through both barriers When penetrates changes in PM Highly controlled process o Egg surrounded by zona pellucida surrounded by cumulus Cumulus various ovary cells Zona pellucida glycoproteins (like vitelline envelope) Testis and sperm o Elongated cell with 3 regions Head Nucleus and acrosome Forms cap over nucleus Packed with enzymatic proteins Midpiece Packed with mitochondria Tail o Produced in seminiferous tubules Mature sperms released in the lumen Tissue between tubules Ledig cells Secrete testosterone Contain Sertoli cells nurture developing sperms Secrete hormone called inhibin Fertilization o Several distinct steps Species specificity in mutual recognition of gametes Activation of sperm Fusion of sperm and egg plasma membrane Blocking of entry by additional sperm Egg and sperm nuclei fuse diploid zygote o Species-specific recognition mechanisms located in zona pellucida Activation of the sperm and the acrosomal reaction o Sperm of same species makes contact with egg Biochemical in jelly trigger acrosomal reaction Breakdown of membrane covering acrosome Enzymes released Enzymes bore hole in jelly and vitelline envelope head Polymerized actin elongates & carries bindin Egg membrane has bindin receptors o 2 types of actions prevent entry of more sperm Fast block to polyspermy few seconds Slow Sodium ions flood eggchange electrical property block to polyspermy 30 seconds Bindin receptors removed Vitelline envelope is hardened Hormonal regulation of reproduction 2 important hormones released by hypothalamus o Gonadotropin-releasing hormone (GnRH) o Prolaction releasing hormone o Stimulate anterior pituitary Follicle-stimulating hormone (FSH) Leutinizing hormone (LH) Women at puberty hypothalamus increases release of GnRH o Increase of FSH and LH targets ovarian tissue Produces estrogen and progesterone Development of sexual characters o Estrogen has time-dependent effect on ovarian cycle Menstruation is beginning of cycle 1st 12 days FSH & LH negative feedback by estrogen on 12th day positive feedback increase in LH & FSH Stimulates follicle cells to make corpus luteum Endocrine gland estrogen & progesterone o Progesterone prepares uterus for pregnancy Development of endometrium on uterine lining If no fertilization corpus lutem degrades on 26th day Endometrium degenerates without progesterone Discarded through menstruation o Decrease in estrogen & progesterone releases neg. control o Cycle starts again! Hormonal control of male reproductive system o Spermatogenesis and development of sexual characters Depends on secretion of testosterone FSH and LH control testosterone release by Ledig cells Under the control of GnRH Testosterone and inhibin exert negative feedback inhibition of hypothalamus and pituitary Evolutionary perspectives on reproductive systems in vertebrates Their evolution is related to migration from water to land Behavioral and structural adaptations promotes proximity of eggs and sperm enhances chances for fertilization Amphibians proximity through copulation Reptiles and birds amniote egg with sufficient food and water o Permeable to air but not water o Fertilization before formation of shell internal fertilization ALL MAMMALS PRACTICE INTERNAL FERTILIZATION Homology between GnRH and mating alpha-factor in yeast o Can bind to mammalian receptor for GnRH Needs higher concentrations for same effect Aging, senescence and death Life span of organisms and cells Potential life span genetically determined o Actual organism-environment interactions Correlaition between life span, length of gestation and puberty age Life span and total number of cell doublings o Related to number of possible divisions by the cells Fixed number of divisions for each cell Positively correlated with life span Cell stop dividing telomeres not long enough o Reduced by 50 bp at every division Programmed cell death (PCD) or apoptosis Cellular/molecular basis of cell death o Cell triggers activation of specific genes products kill cell o Time and place of PCD genetically programmed Nucleus condenses Nuclear DNA is cut by DNAse between nucleosomes Ladder pattern of DNA fragments gel electrophoresis PCD in normal development Several genes regulate ced3 & ced4 promote, ced9 inhibits PCD in animal development o Major role in limb development carves limb bud Chiseling generates variation o Pattern of selective PCD determined by mesoderm PCD in plant development o Major role in development of plant body Xylem, leaf trichomes, egg development o Role in stress response Cell with pathogen die through PCD with pathogen Rest of plant is saved Different from necrosis o PCD PM is intact o Necrosis PM deteriorates cell lysis Patterns of senescence in plants Cumulative effects of deteriorative processes leads to death o Genetically programmed and developmentally regulated Whole plant senescence o Whole plant, except seed, dies Senescence of lateral organs in trees o Lateral organs such as leaves die Senescence of above-ground parts only o Shoot dies every year but underground parts survive When does senescence start o At whole organism onset of sexual maturity o Rate of senescence related to life span of organism or organ o In leafs Total protein and chlorophyll content starts declining Membranes become leaky With increase in lipid peroxidation products Reversal of foliar senescence o Leaf senescence also triggered by lead maturity Reverse senescence by removing upper part of plant Hormonal and anti-oxidant regulation of leaf senescence o Membrane deterioration mediated by anti-oxidants Ethanol, vitamin E, isobenzofuran o Cellular levels of some hormones and anti-oxidant defenses of the plant can regulate rate and intensity of leaf senescence Sensing the environment – Nervous System Nervous system – Introduction 2 majors parts central and peripheral nervous system o CNS brain & spinal chord o PNS nerves Cells in nervous system highly excitable & conduct electricity o Neurons Information transfer and storage o Glial Out-number neurons 5:1 Schwann cells Wrap body many times around neuron in PNS Form myelin sheet Insulate axon & increase conduction speed Maintenance and repair of damaged neurons Digests debris & guides re-growth Astrocytes Star-shaped body in CNS Multiples contacts with neurons, myelinated part of axons, blood capillaries & ependymal cells Gap-junctions Transport nutrients & regulate extracellular space Remove debri from damage & guide development Oligodendrocytes Myelin sheet around neuronal axons of CNS Can for myelin around several nearby neurons Microglia Smallest glial cells located in CNS Remove debris from trauma or disease Ependymal cells Line ventricles of CNS Cavities filled with cerebrospinal fluid Cilia on surface circulate cerebrospinal fluid Processes that makes connections with Astrocytes Cell types in nervous system: Neurons Wide main body called soma rich in cell contents Individual branches dendrites o Make contacts with other neurons synapses Long process coming out of soma axon o Region of connection axon hillock Information is integrated o Axon is myelinated narrow bare part node of Ranvier Each sheath is internode Types of neurons o Purkinje cell fewer but highly branched dendrites o Many other types, see page 6 of lecture Structural classification position & number of dendrites + location of integrating center o Multipolar many dendrites and 1 long axon (myelin) o Bipolar single dendrite and 1 long axon (no myelin) o Unipolar single long myelinated axon o Pseudo-unipolar 2 axons Functional classes o Motor efferent take information to muscles o Sensory afferent take information from sensory organs o Interneurons connects previous 2 to CNS Action potential: generation and perpetuation Unequal distribution of charges across PM -70 mV o Inside more negative than outside Entry of positive ions membrane depolarization More negative than -70 mV hyperpolarization Stimuli which depolarize by causing specific ion channel to open o Potential below threshold graded potentials o Potential above threshold active potential Only voltage-gated ion channels Wave action of membrane depolarization o Local potential arises to maintain magnitude of action o Myelination of axon increases speed Nodes of Ranvier 1mm apart potential jumps The synapses and the neurotransmitters Synapse composed of presynaptic and postsynaptic cells and synaptic cleft Electrical synapse electrical signal passes as electrical signal Chemical synapse release of neurotransmitter Neurotransmitters o Small molecular weight molecule o Synthesized in neurons o Release triggered by action potential at distal end Neuromuscular junction Neurotransmitter released is acetylcholine Neuron originates from CNS At distal end axons divides into several branches o End in terminal button Forms a synapse with single muscle fiber Motor endplate Presynaptic voltage-gated Ca channels & Ach vesicles Postsynaptic Ach receptors Action potential opens Na channels depolarization Ca channels open Ca released Fusion of Ach vesicles with PM Ach released in cleft Binds to receptors activation open ion-gated channels Depolarization leading to muscle action Ach released and degraded in cleft presynaptic recycles Invertebrate nervous system Cnidarians have simplest NS o Neurons are not specialized impulses in all directions In evolution network rearranged in pathway with defined flow o Ganglion cell bodies fuse together with synapses Integrating centers of varying complexity Bilaterally symmetrical animals complex groupings brain o Ganglia are arranged in series o Cephalization locating sense organs & integrating centers at anterior end of the body Vertebrate nervous system 3 divisions of nervous system o Integrating center brain & spinal chord CNS o Smatosensory division sensory neurons o Motor division motor neurons Brains o Narrow neural groove from neurula stage results in brain o Protected by cranium & meninges Dura mater, arachnoid mater & pia mater o Forebrain, midbrain and hindbrain Hindbrain leads to spinal chord Peripheral nervous system o Cranial and spinal nerves o 2 main divisions Somatic motor nervous system voluntary control Autonomic nervous system no conscious control Sympathetic Active in stressful conditions Regulates blood flow and pressure Parasympathetic Active in periods of rest and calm Redirects energy Enteric Independent gastro-intestinal tact, etc. Structure of vertebrate nerve o Several bundles of axons bundle = fascicle Individual neuron surrounded by endoneurium Fascicle surrounded by perineurium Nerve surrounded by epineurium o Forms synapse at both ends Structure of a reflex arc Receptor stimulus perceived Sensory pathway sensory neurons take info to CNS Integrating center decision making Motor pathway info from integrating sensor to muscles Effector organ organ where the action is taken 4/12/2010 8:50:00 PM 4/12/2010 8:50:00 PM 4/12/2010 8:50:00 PM