Book pp. 928–930 · asked 6 times in NTRUHS papers

In one breath

CO₂ travels from the tissues to the lungs in three forms: about 70% as bicarbonate, 23% bound to proteins as carbamino compounds, and 7% dissolved. Carbonic anhydrase in the red cells makes the bicarbonate; the chloride shift swaps it out into plasma. Because deoxygenated Hb holds more CO₂ and H⁺, releasing O₂ in the tissues helps blood take up CO₂, and taking up O₂ in the lungs helps blood let CO₂ go. That second half is the Haldane effect.

Builds on: Oxygen transport · Oxygen–hemoglobin dissociation curve · Leads to: Chemical control of breathing · Asphyxia, hypercapnia and hypocapnia

CO₂ is made mainly by aerobic metabolism of glucose (and when carbohydrate is turned into fat). Venous blood carries it to the lungs, where it is breathed out.

FormShareWhere
Bicarbonate (HCO₃⁻)~70%Formed in red cells, mostly carried in plasma
Carbamino compounds~23%Mainly carbamino-Hb in red cells; a little with plasma proteins
Dissolved~7%Plasma

Dissolved form in plasma

  • Dissolved CO₂ follows Henry’s law (see Oxygen transport): it is proportional to PCO₂.
  • CO₂ is about 20 times more soluble than O₂: its solubility is 0.06 mL/dL per mmHg.
  • In venous blood: 46 mmHg × 0.06 ≈ 2.8 mL/dL dissolved.
  • Plasma has no carbonic anhydrase, so little carbonic acid forms there. A small amount of CO₂ binds plasma proteins as carbamino-protein.

As bicarbonate ions

  • Most CO₂ diffuses straight into the red cells, which are rich in carbonic anhydrase (CA):

    CO₂ + H₂O ⇌ (CA) ⇌ H₂CO₃ ⇌ H⁺ + HCO₃⁻

  • What happens to the H⁺: the red-cell membrane barely lets H⁺ through, so it stays inside and is buffered by Hb:

    H⁺ + HbO₂⁻ ⇌ HHb + O₂

    • Binding H⁺ lowers Hb’s O₂ affinity (the Bohr effect), so O₂ is released to the tissue, and the H⁺ is mopped up, so more CO₂ can be converted. The two gases help each other.
    • In the lungs everything runs backwards: oxygenation of Hb drives CO₂ back out.
  • The chemistry happens in the red cells, but most of the bicarbonate then rides in the plasma, after the chloride shift.

Chloride shift

  1. HCO₃⁻ builds up in the red cell and moves out into plasma.
  2. To keep the cell electrically neutral, Cl⁻ moves in in exchange.
  3. The exchange is carried by the anion exchanger 1 (AE1) in the red-cell membrane, formerly called the band 3 protein.
  4. This Cl⁻-for-HCO₃⁻ swap is the chloride shift, also called the Hamburger shift after the scientist who described it.
  5. In the lungs the shift reverses: HCO₃⁻ goes back into the cell, and Cl⁻ comes out.

Applied: why venous hematocrit is higher

Venous red cells contain about 20% more Cl⁻ than arterial ones. Cl⁻ is osmotically active, so the cells take in water and swell. That makes the venous hematocrit about 3% higher than the arterial one.

Draw it: the chloride shift

Draw a red cell beside a tissue capillary. CO₂ enters, then CO₂ + H₂O → (CA) → H₂CO₃ → H⁺ + HCO₃⁻. Draw H⁺ binding to Hb (with O₂ leaving to the tissue), HCO₃⁻ leaving through AE1 (band 3), and Cl⁻ entering through the same exchanger. Add a small “H₂O in, cell swells” arrow.

As carbamino compound

  • CO₂ binds the free amino (–NH₂) groups of Hb directly, forming carbaminohemoglobin:

    CO₂ + Hb–NH₂ ⇌ Hb–NH–COOH

  • Deoxy-Hb binds more CO₂ than oxy-Hb, so unloading O₂ in the tissues boosts this route (part of the Haldane effect).

  • About 23% of CO₂ travels this way.

CO₂ dissociation curve

0204060800204060PCO₂ (mmHg)CO₂ content (mL/dL)avPO₂ 40 (deoxygenated)PO₂ 100 (oxygenated)
The CO₂ dissociation curve: nearly straight, and lower when blood is oxygenated (the Haldane effect).
  • A plot of blood CO₂ content against PCO₂, made like the O₂ curve.
FeatureCO₂ dissociation curveO₂–Hb dissociation curve
ShapeAlmost a straight line over the working rangeSigmoid
Over the working rangeSteep and linear between PCO₂ 40 and 46 mmHgMostly flat between PO₂ 40 and 100 mmHg
SaturationNo saturation point in the physiological rangeSaturates near 100 mmHg
Volume carriedBlood holds much more CO₂ than O₂~20 mL O₂/dL
  • Because the curve is steep, a small change in PCO₂ loads or unloads a lot of CO₂.
  • Worth knowing, though not in your pages: your chapter makes the comparison without giving figures. The standard values are a total CO₂ content of about 48 mL/dL in arterial blood and about 52 mL/dL in venous blood.
  • Effect of PO₂: at higher PO₂ the whole curve moves down and to the right. At any given PCO₂, blood holds more CO₂ when its PO₂ is low. This inverse relationship is the Haldane effect.

Haldane effect

  • Definition: oxygenation of Hb lowers its affinity for CO₂ (and H⁺), so taking up O₂ in the lungs drives CO₂ out of the blood. The reverse happens in the tissues: releasing O₂ lets blood take up more CO₂.
  • Why it happens:
    1. Oxy-Hb is a stronger acid and binds fewer H⁺, so H⁺ is released, combines with HCO₃⁻ and is breathed out as CO₂.
    2. Oxy-Hb forms fewer carbamino compounds.
  • Named after John Scott Haldane (1860–1936).
  • Worth knowing, though not in your pages: the Haldane effect accounts for close to half of the CO₂ unloaded in the lungs.

Bohr effect

  • Definition: a rise in PCO₂ or H⁺ (and temperature) lowers Hb’s affinity for O₂, which shifts the O₂–Hb curve right and raises P50.
  • In the tissues, CO₂ enters the blood, the pH falls, and Hb gives up more O₂. Tissue oxygenation improves exactly where metabolism is high.
  • Mechanism: mainly the drop in pH, since deoxy-Hb binds H⁺ more avidly than oxy-Hb; a rise in temperature adds to it.
  • Named after Christian Bohr (1855–1911), a Danish physiologist.
Bohr effectHaldane effect
What changesCO₂ / H⁺O₂
What it affectsHb’s affinity for O₂Blood’s affinity for CO₂
Curve it shiftsO₂–Hb curve → rightCO₂ curve → down and right
Where it matters mostTissues: helps unload O₂Lungs: helps unload CO₂

Don't copy your book's one-line summary

The book’s Key Concepts pair the effects confusingly (“Haldane helps loading of O₂… Bohr helps loading CO₂”). Remember it this way instead:

  • Bohr is CO₂ pushing O₂ off Hb in the tissues.
  • Haldane is O₂ pushing CO₂ off blood in the lungs.
  • Each gas helps the other leave.

Exam-answer skeleton: "Describe the transport of CO₂ in blood" (long essay)

  1. Where CO₂ comes from, and the three forms with their percentages.
  2. The dissolved form: Henry’s law, solubility 20× O₂, 46 × 0.06; no CA in plasma.
  3. Bicarbonate: the CA reaction, H⁺ buffered by Hb, then the chloride shift (with a diagram) and why venous Hct is higher.
  4. Carbamino-Hb: the reaction; deoxy-Hb binds more.
  5. The CO₂ dissociation curve: its shape compared with the O₂ curve (with a diagram).
  6. The Haldane effect and the Bohr effect, with the comparison table.
  7. The reverse events in the lungs.

Asked in exams