Topic 5 (part 2) - In person Lecture

Define respiration

  • process that provides energy for life - a sequence of events that results in the exchange of O2 and CO2 between the external environment and the animal + its mitochondria within its cells

what is respiration’s purpose in organisms?

  • purpose of respiration is gas exchange of O2 and CO2 → receives oxygen from the environment and removes carbon dioxide from blood

Fick’s law of diffusion:

k - diffusion constant

  • intrinsic property of the gas that relates to how well it can move through the membrane

  • independent of specific pressure difference or thickness of the membrane

  • depends on solubility of the gas and temperature

A - area for gas exchange

(P2-P1) - difference in partial pressure of gas on either side of barrier to diffusion

D - distance → thickness of barrier to diffusion

How to increase/decrease the rate of diffusion based on the parameters in the equation

increasing:

  • diffusion constant of gas is large/high

  • surface area of membrane is large

  • difference in partial membrane of gas is high on either side of membrane

  • diffusion distance across membrane is small

decreasing:

  • what diffusion constant of gas is small/low

  • surface area of membrane is small

  • difference in partial membrane of gas is low on either side of membrane

  • diffusion distance across the membrane is large

Graham’s law:

  • diffusion constant looks at intrinsic gas properties of solubility and molecular weight → based on Graham’s Law

MW - molecular weight

\left(MW\right)^{0.5}=\sqrt{\left(MW\right)}

rate of diffusion for gases can also increase when:

  • solubility of a gas in medium is high

  • molecular weight of a gas decreases

  • liquids - solubility is a more significant factor than molecular weight / air - molecular weight is the more significant factor than solubility

what is ventilation

  • the active movement of respiratory medium (air/water) across respiratory surface

What are the three types of ventilation?

  • non-directional - medium flow past the respiratory surface in an unpredictable pattern

    • animals that use cutaneous respiration (i.e. earthworms, frogs, lungless salamanders) or wave their external gills directly into the environment medium → are examples that follow this ventilation pattern

  • unidirectional - medium moves through respiratory system in one direction - passes over gas exchanges before exiting through a separate pathway

    • seen in fishes with internal gills (teleost/elasmobranch gills) + avian lungs

  • tidal - medium moves in and out of respiratory chamber in a back and forth movement

    • seen with lungs of mammals, insects, most reptilians (non avian - except crocodilians), air breathing amphibians, air breathing fish

Non-directional ventilation:

  • partial pressure of oxygen in blood leaving the gas exchanger can approach the partial pressure of oxygen in the medium - if the medium is very well mixed

  • anything that increases diffusion distance - decreases the oxygen exchange efficiency and reduces the partial pressure of oxygen in the blood leaving the gas exchanger

Tidal ventilation:

  • when an animal breathes in - incoming fresh medium mixes with stale medium left from previous breath in respiratory cavity

    • partial pressure of oxygen in respiratory cavity is lower than that of the external environment

  • oxygen diffuses into blood flowing along the gas exchange surface

  • partial pressure of oxygen in the blood exits the gas exchange surface in an organism - will be approximately in equilibrium with the exhaled medium (if diffusion distance across respiratory surface is small)

Uni-directional ventilation:

  • blood can flow in two ways relative to the flow of the medium

    • countercurrent flow - opposite direction along the gas exchange surface

    • crosscurrent flow - at a perpendicular direction

countercurrent flow - medium starts with high partial pressure of oxygen and decreases steadily as it gives up oxygen along the exchange surface

  • blood starts with low partial pressure of oxygen and steadily increases as it picks up oxygen while flowing in opposite direction to medium

  • medium always has a higher partial pressure of oxygen than blood flowing next to it - creates a small but continuous diffusion gradient across the entire exchange surface (key to efficiency)

crosscurrent flow - medium starts with high partial pressure of oxygen and decreases steadily as it give up oxygen along exchange surface

  • blood starts with low partial pressure of oxygen and steadily increases as it picks up oxygen from different parts of the medium - blood’s partial pressure of oxygen can rise to a level higher than the final partial pressure of the medium

Which of these flow systems is more efficient at gas exchange

  • countercurrent exchange is more efficient at gas exchange - maintains a consistent diffusion/thermal gradient along the entire exchange surface rather than allowing it to drop quickly like in crosscurrent flow

Boyle’s Law - states that in a closed space pressure and volume are inversely related

  • meaning if volume decreases = pressure increases - vice versa

Fish’s gills

  • elasmobranchs - most species of sharks, skates and rays → used buccal pump for gill ventilation

    • mouth opens → buccal cavity expands in volume → decreases pressure inside the cavity

    • water rushes/is sucked into buccal cavity → occurs through the mouth and spiracles (nostril-like structures on top of head)

    • mouth and spiracles close - muscles surrounding buccal cavity contract → decreases volume of cavity + increase pressure inside → forces water past gills and through external gill slits

    • buccal pump → a dual-pumping mechanism in which buccal cavity alternates suction/force phases → to move water unidirectionally over the gills (even if the animal is stationary)

  • gills in teleost (bony) fish - located in opercular cavities

    • are protected by flap-like operculum (provides better protection of gills compared to more exposed gills seen in elasmobranchs)

    • 4 gill arches in each opercular cavity - provide structural support for two rows of gill filaments (i.e. primary lamellae) that project from each gill arch in a v-shape

    • each filament is covered with rows of interdigitated folds called secondary lamellae → perpendicular to the filament

    • secondary lamellae are thin-walled + highly vascularized → acts as a main respiratory surface

    • uses a buccal-opercular pump for gill ventilation - requires buccal (mouth) and opercular (gill cover) action in tandem to drive water in a continuous unidirectional flow across internal gills

Ventilatory cycle of teleosts

  • mouth opens / opercular valve closed - buccal cavity expanded + opercular cavity expands

    • expansion creates negative pressure - drawing water into the mouth from the outside

  • mouth closes / opercular valve closed - buccal cavity is compressed + opercular cavity is expanded

    • negative pressure in opercular cavity helps move water from buccal cavity to gills

  • mouth closed / opercular valve opens - buccal cavity is compressed + opercular cavity compresses

    • compression of both buccal cavity + opercular cavity force water over the gills and out

  • mouth opens / opercular valve opens - buccal cavity expands + opercular cavity compressed

gas exchange is effective at fish gills:

  • lamellae have a very thin membrane → small diffusion distance

  • lamellae provide very large surface area for gills

  • large partial pressure gradient of gas between water column and capillaries inside lamellae

fish gills - use a countercurrent exchange system → where blood flow through capillaries in secondary lamellae is in the opposite direction to water passing through gills

  • oxygen extraction from water can be high as 70-80%

  • counter current exchange system is a good way to compensate with an oxygen deprived environment → but air holds significantly more oxygen than water

why crosscurrent is less efficient:

  • inefficiency in oxygen extraction - if blood + water flowed in the same direction it would reach to equilibrium + oxygen diffusion would stop → lower oxygen saturation levels in the blood - insufficient for metabolic needs of most fish

  • low oxygen concentration - water contains much less oxygen than air - in order to survive fish need an extremely efficient method to extract the maximum amount of oxygen possible

Ram ventilation

  • method of respiration where fish (i.e. teleost fish - tuna, salmon, flounders, cod, herring + tiger shark) swim forwards with their mouths open to force water across their gills, facilitating oxygen uptake without active pumping

    • some sharks (tiger sharks) can breathe by alternating between buccal pumping → swimming at low speeds + ram ventilation → swimming at higher speeds

    • some sharks (i.e. great white, whale shark, hammerhead + mako) don’t have buccal muscles → have to be ram ventilators - need to keep moving forward in order to breathe (if they stop they will drown)

advantages:

  • fish can prioritize more energy into their swimming muscles instead of their buccal or opercular muscles → enhances their speed

  • increase flow rate of water over gills as fish swims faster → increased oxygen uptake to meet higher metabolic needs of fish

disadvantages:

  • some obligate ram ventilators must swim continuously to breath for survival

  • opening mouth results in increased drag on fish → affects locomotion - increases the resistance against the water and slows the fish down (drag is minimized if they have a streamlined body - designed to minimize water resistance [drag])

streamline body

Air breathing fish

mudskippers - spend more time on land than in water - some species spend 90% of their time out of the water

  • absorbs oxygen through their skin (cutaneous respiration) + through the lining of their mouth + throat → both contain a high density of blood capillaries

  • modified gill chambers - helps trap pockets of air + water while on land → prevents the gills from collapsing and allows for gas exchange

electrical eels

  • have extensive folds in mouth - highly vascularized for substantial aerial gas exchange (aerial gas exchange → exchange oxygen + carbon dioxide directly with the atmosphere)

highly vascularized → has an exceptionally high density or concentration of blood vessels (i.e. - arteries, veins, capillaries)

armored catfish

  • highly vascularized stomach that is used for gas exchange after gulping air

  • can survive 30 hours out of the water if enough oxygen is stored in their stomachs

bichirs

  • uses a pair of lungs that are highly vascularized for gas exchange

  • are facultative air-breathers - possesses both gills + specialized organ to breathe air (typically only does one when necessary)

  • can access surface air to breathe when the water they inhabit is poorly oxygenated

lungfish - most developed air-breathing organ of any fish

  • have complex highly vascularized lungs → covered in folds + pockets that increase their surface area

  • most lungfish species have two symmetrical lungs (except Australian lungfish → only has one lungfish)

Respiration in Insects

tracheal system - a complex network of tubules that insects use in order to breathe

  • large tracheae is connect to spiracles

    • spiracles - openings on the surface along the abdomen where oxygen enters and CO2 leaves

    • spiracles can be closed using valves to reduce water loss → allows insects to live in dry areas

    • rise in CO2 in the tracheal system causes spiracles to open

  • air sacs are reservoirs (pushes air through tracheal tubes when contracted)

  • tracheal tubes + tracheoles are tubes that carry air throughout the body directly to cells and tissues (i.e. muscles)

    • gases are moved largely by muscle contractions in their abdomen and around their spiracles → no blood flow is needed to transport gases

    • *air is not circulated by the insects blood - respiratory + circulatory systems are separate

Respiration in Amphibians + non-avian reptiles

frog lungs are not as efficient as fish gills:

  • new air that is inhaled mixes with the old air that is already in the lung (note: tidal ventilation is not as efficient in gas exchange compared unidirectional ventilation)

  • the lungs have a relatively smaller surface area

  • oxygen levels are much higher in the air (about 21% oxygen level) than in water (1% oxygen level) → lungs do not have to be as efficient as gills

why non-avian reptiles have more efficient lungs than frogs

  • non-avian reptiles are generally more active than amphibians - require more oxygen

  • lungs of non-avian reptiles contain a larger surface area compared to the lungs of amphibians

why non-avian reptiles cannot use cutaneous respiration like frogs

  • cannot rely on their skin for respiration (like amphibians) due to dry scaly skin → water tight to avoid water loss

Human Respiratory system

identify parts of the human respiratory system

purposes of the C-shaped hyaline cartilaginous rings - trachea:

  • keeps the trachea open during breathing (prevents collapsing) so oxygen can reach the alveoli

  • rings are incomplete (c-shaped) instead of complete (o-shaped) so that the esophagus can expand during peristalsis as food or liquid moves through it

peristalsis - series of involuntary, wave-like muscle contractions that propel food, fluids, waste through the digestive tract

how gas exchange is very effective at the alveoli based on Fick’s law

  • alveoli + capillaries are one cell layer thick → have a small diffusion distance

  • alveoli provide very large surface area for lungs

  • large partial pressure gradient of gas between inside of alveoli and capillaries

Respiration in birds

Identify: trachea, primary bronchus, ventrobronchus, dorsobronchus, parabronchi, anterior air sacs, posterior air sacs (do not need to know the name of the air sacs, posterior and anterior is good)

  • unidirectional air flow through respiratory system

  • lungs are almost completely rigid + undergoes little change in volume during breathing

  • birds require two cycles of breathing (inhalation + exhalation) to move air through its respiratory system

    • they inhale by expanding the volume of its chest because of contracting rib (intercostal) muscles + muscles attached to the sternum (breastbone)

      • *birds do not have a diaphragm like mammals do

      • expansion of chest volume causes increase in volume of air sacs while pressure within them decreases → causes air to rush in along its partial pressure gradient

pathways of first + second of inhalation and exhalation:

  • first inhalation

    • anterior + posterior air sacs expand

    • air travels through trachea → splits into left + right primary bronchi

    • air then travels down each primary bronchus and primarily into posterior (rear) air sacs

  • first exhalation

    • chest compresses while anterior + posterior air sacs compresses → volume in air sacs decrease + pressure increases

    • air moves along partial pressure gradient from posterior sacs into lungs where gas exchange occurs

  • second inhalation

    • anterior + posterior air sacs expands

    • air moves from lungs into anterior sacs

  • second exhalation

    • anterior + posterior air sacs compress

    • air moves from anterior sacs back into trachea and expelled from the system

how avian respiratory system is more efficient at gas exchange over mammalian respiratory system:

  • unidirectional nature - higher oxygen content that can diffuse into blood and reach tissues

  • wall of lungs cannot collapse in on themselves → does not require residual air since avian lungs are more rigid

    • residual air - approx 1-1.2 L of air that permanently remains in the lungs even after a maximum forced exhalation - prevents the alveoli (air sacs) from collapsing + keeps it inflated

    • mechanism eliminates the need to mix fresh air with old air in lungs → ensures a constant supply of oxygenated air across the respiratory surface

      • mixing of air reduces the partial pressure of oxygen and increases the partial pressure of carbon dioxide within the alveoli → reduces efficiency of gas exchange in accordance to Fick’s law

  • air capillaries in walls of parabronchial system having much larger overall surface area than alveoli found in mammalian respiratory system