1 RESPIRATORY SYSTEMS Functions: 1º gas exchange: O2/CO2 water/acid-base balance 2º (air-breathers) vocalizations, thermal balance Basic requirements: -high surface area -short diffusion distance for exchange -gas-exchange region highly vascularized BREATHING MEDIUM - RELATED FACTORS medium sets energy costs to breathe -water high mass, low O2 content → expensive -air low mass, high O2 content → cheap breathing cost sets ventilation mode -water: unidirectional (mostly) → GILLS (AND SKIN) -air: tidal (birds unidirectional) → LUNGS 2 VENTILATORY ORGANS: OLD OR NEW? What are basic requirements for gas exchange? -high surface area -short diffusion distance for exchange -gas-exchange surface highly vascularized Adapt what already exists? -skin advantages: next to medium, vascularized, thin disadvantages: protection function in conflict, SA low -gut: advantages: access to medium (pharyngeal region), vascularized, SA can be increased New structures based on gut -extend portion of pharynx wall into medium → GILLS -develop internal blind sac from pharynx wall → LUNGS 3 RESPIRATION IN FISH - GILLS -breathing medium: water - dense, low O2 content -high metabolic cost to breathe -unidirectional flow - most efficient -gas exchange membrane extended into breathing medium FISH GILL ORGANIZATION Kardong f 11-4 4 TELEOST GILL VENTILATION PUMP CYCLE -UNIDIRECTIONAL FLOW OVER GILLS Eckert f 14-31 TELEOST GILL STRUCTURE gill bar bone or cartilage Eckert f 14-33 5 HISTOLOGY OF GILL LAMELLAE 2º lamellae attached to filaments cell types: epithelial cells endothelial cells = pillar cells chloride cells (ionophores) mucus cells PILLAR CELLS (=ENDOTHELIAL CELLS) DEFINE BLOOD SPACES IN 2º LAMELLAE Patt and Patt (1969) f 8-1 6 CORROSION CASTING OF BLOOD SPACES IN 2º LAMELLAE Eckert f 14-34 BLOOD FLOW THROUGH TELEOST GILL - COUNTERCURRENT EXCHANGE Eckert f 14-35 7 BLOOD AND WATER FLOW IN OPPOSITE DIRECTIONS PAST 2º LAMELLAE = COUNTERCURRENT EXCHANGE Patt and Patt (1969) f 8-2 LAMPREY GILLS LINE GILL POUCHES -mouth attached to food -blocks water intake through mouth -water flows into and out of each pouch via gill openings Eckert f 14-32 8 LAMPREY GILL POUCHES dorso-ventral pumping muscles cartilagenous branchial arch pharynx Kardong f 11-14 external surface gill pouch opening 9 LUNG VENTILATION IN VERTEBRATES ADVANTAGES OF AIR: cheap cost to breathe: high O2 content, low mass PROBLEM: can't extend gas exchange membrane into medium -still need aqueous layer on membrane surface -gases dissolve in aqueous layer -moisture loss damages membrane TRENDS IN AIR-BREATHING -internalize gas exchange organ: develop blind sac from gut -increase internal surface area for gas exchange: septation ↑ -decrease blood-air barrier thickness: diffusion distance ↓ 10 GAS EXCHANGE AREA INCREASES WITH MORE SEPTATION Hildebrand f 13-8 LUNG VENTILATION no ribs: positive pressure ventilation amphibians ribs: negative pressure ventilation reptiles, birds, ribs + diaphragm: mammals 11 AMPHIBIAN RESPIRATION ALL MODES GILLS: ALL LARVAL AND SOME ADULT FORMS SKIN: ALL AQUATIC AND MOST SEMI-AQUATIC FORMS LUNGS: SEMI-AQUATIC AND TERRESTRIAL FORMS LUNGS: TIDAL VENTILATION: POSITIVE-PRESSURE SYSTEM -BUCCAL PUMP - JAW MUSCLES (NO RIBS) STRUCTURE: LUNG SEPTATION MINIMAL TRACHEA, BRONCHI SHORT CILIATED EPITHELIUM, GOBLET CELLS SEPTAL WALLS THICK BLOOD-AIR BARRIER THICK POSITIVE-PRESSURE VENTILATION INSPIRATION: nares open, jaw floor drops, air into buccal cavity, nares close, jaw floor rises, air into lungs EXPIRATION: nares open, elastic recoil in lungs pushes air out Eckert f 14-27 12 BLOOD-AIR BARRIER IN FROG LUNG SEPTUM Patt and Patt f 8-14 REPTILIAN RESPIRATION RIBS APPEAR → NEGATIVE PRESSURE VENTILATION LUNGS PAIRED (snakes, lizards may have one lung reduced or absent) TRACHEA, 1º BRONCHI cartilage reinforced, ciliated with goblet cells SEPTATION MORE COMPLEX FAVEOLI - major and minor subdivisions -gas exchange in septal walls 13 LIZARD LUNG SHOWING FAVEOLAR DIVISIONS Gans and Gaunt (1998) f 1-15 FAVEOLAR LUNG STRUCTURE F FAVEOLUS SF SUBFAVEOLUS gas exchange in septal walls Gans and Gaunt (1998) f 1-6 14 SNAKES: USE ONE LUNG, OTHER REDUCED OR ABSENT gas exchange region - anterior - vascular - faveolar septation saccular region - no gas exchange - no vascularization - no septation Gans and Gaunt f 2-3 FAVEOLAE IN GAS EXCHANGE REGION OF SNAKE LUNG Gans and Gaunt f 2-4 15 BIRD LUNG HIGH EFFICIENCY - supports flight metabolic requirements UNIDIRECTIONAL VENTILATION cross-current gas exchange between blood and air capillaries LONG CONDUCTING AIRWAYS - HIGH "DEAD" VOLUME Eckert f 14-9 16 BIRD LUNG AND AIR SACS trachea lung cranial air sac caudal air sacs Pough et al f 17-15 AIRWAYS IN CROSS-SECTION OF BIRD LUNG: 1º BRONCHUS→ DORSOBRONCHUS → PARABRONCHI → VENTROBRONCHUS direction of air flow Hildebrand f 13-12 17 UNIDIRECTIONAL AIR FLOW IN BIRD LUNG Eckert f 14-26 ANATOMY OF BIRD LUNG Kardong f 11-35 18 PARABRONCHUS STRUCTURE Eckert f 14-25 PARABRONCHI WITH INFUNDIBULI IN WALLS Farner and King (1972) p 294 CROSSCURRENT GAS EXCHANGE IN BIRD LUNG -most efficient in vertebrates Hildebrand f 13-11 19 COMPARISON OF AIR FLOW AT GAS EXCHANGE MEMBRANE IN MAMMALIAN AND BIRD LUNGS TIDAL Kardong f 11-35 UNIDIRECTIONAL