Lecture 15 Respiratory Systems and Gas Bladders

Kardong Chapter 11, Hildebrand Chapter 13

Respiratory Systems
Organs for external respiration (i.e., gas exchange with the environment) are required by large, active animals like vertebrates. Gases include O2 and CO2.

Requirements of external respiratory surfaces: Large surface area. Thin barrier between blood and air or water. Flow or exchange of air or water. Favorable diffusion gradient between blood and air or water.

The Early Development of Gills

Gill openings develop in the pharynx from inside out and from front to back. Each branchial arch between these openings contains an element of the splanchnocranium (not shown) and a branch of the ventral aorta.

KK 10.22, H&G 13.1

Gills KK 11.4, H&G. 13.2

Lamprey gills are called pouched gills and the 7 openings are almost tubular. Sharks have 5 gill openings covered by septa. There is a half-gill or hemibranch on the hyoid arch, plus 4 holobranchs. The opening ahead of the hyoid arch is the spiracle.

Teleosts have 4 branchial arches without septa, covered by the operculum.

External Gills KK 11.4, H&G 13.5

External gills are larval structures found in some Osteichthyes, including Sarcopterygii, and Amphibia (remember Necturus?).

In fish, external gills develop before pharyngeal gills and are an adaptation to low O2 environments. (Larval fish are small enough that under most conditions they do not need gills.) Tetrapods, including Amphibia, never have functional pharyngeal gills.

Holobranch of a shark.
KK 11.17, H&G 13.2

Shark gills are protected by the gill septa separating the two halves of the holobranch.

Teleost Gills KK 11.19, H&G 13.3

Teleost gills are free, in the sense that they do not have a gill septum. They are protected by the operculum instead. Note the direction of blood flow in the secondary lamellae, which are like fins on a radiator.

The direction of water flow across the gills is opposite to the direction of the blood flow within the gill lamellae.

That the water and blood flow in opposite directions is important; it maintains a constant +ve gradient for gain of O2 or loss of CO2. This is called countercurrent gas exchange.

Lungs

Lungs are evaginations of the anterior gut behind the gills. Their surfaces are highly folded and vascularized to maximize gas exchange. This is not true of fishes that have lungs that they do not use (e.g., sturgeon) or fishes that have gas bladders (teleosts). Most fish lungs or gas bladders are dorsal in position, while tetrapod lungfish lungs and tetrapod lungs are more ventral. Why lungs?

KK 11.5, H&G

Gas Bladders KK 11.22, H&G 13.7

Although some teleosts maintain a connection from the gas bladder to the gut (e.g., trout) most lose this connection in development. Either way, the gas content of the bladder is maintained by adding or removing gas from the circulatory system via the red body and the oval body, respectively. Some fish use gas bladders for sound reception and/or production.

Respiratory Systems in Tetrapods

Modern Amphibia external gills (larvae) cutaneous respiration mouth breathing lungs, ventilated by pharynx Ancient Amphibia and Reptilia

KK 11.21, H&G 11.21

mouth breathing lungs, ventilated via aspiration by ribs using intercostal muscles Mammals - lungs ventilated via aspiration by ribs, diaphragm in pleuroperitoneal membrane Birds
- lungs, ventilated by sternum and other

The importance of cutaneous respiration varies from 100% to near 0.

KK 11.6

KK 11.36

Mammalian lungs, as illustrated by the hippopotamus, have a large surface area for respiration because of their complex internal structure of alveoli. In humans this area is about 70 m2.

Avian Lungs
Avian lungs are small and constant in volume, but are associated with hollow air sacs that take up space in the body and lighten it. Extensions of the air sacs penetrate even into the bones, so they are hollow.

Air flow through the lungs is achieved not by expanding the lungs (aspiration) but by action of the sternum and muscles on the air sacs. This efficiently capitalizes on the action of muscles during flight.

Diagram of left lung of a bird.

KK 11.37, H&G 13.12
Air passes through bird lungs in one direction, not tidally, through air capillaries called parabronchi. The blood flows through the lung in the opposite direction. This greatly increases the gas exchange efficiency of bird lungs relative to those of other tetrapods.

KK 11.36, H&G 13.10, 13.13

The tidal flow of air in the lungs of mammals and other tetrapods is relatively inefficient compared to the cross-current flow of air through the lungs of birds. Birds can undergo strenuous exercise (flying) at altitudes where we would pass out standing still.