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00:00:01,940 --> 00:00:05,190
So we've talked a little bit
about the lungs and the tissue,

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00:00:05,190 --> 00:00:08,430
and how there's an interesting
relationship between the two

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00:00:08,430 --> 00:00:12,490
where they're trying to
send little molecules back

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00:00:12,490 --> 00:00:13,010
and forth.

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00:00:13,010 --> 00:00:16,079
The lungs are trying to
send, of course, oxygen out

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00:00:16,079 --> 00:00:17,800
to the tissues.

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00:00:17,800 --> 00:00:19,780
And the tissues are
trying to figure out

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a way to efficiently
send back carbon dioxide.

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00:00:23,200 --> 00:00:25,505
So these are the
core things that

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00:00:25,505 --> 00:00:27,080
are going on between the two.

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00:00:27,080 --> 00:00:29,080
And remember, in terms
of getting oxygen across,

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00:00:29,080 --> 00:00:31,060
there are two major
ways, we said.

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The first one, the easy one
is just dissolved oxygen,

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00:00:35,590 --> 00:00:38,240
dissolved oxygen in
the blood itself.

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But that's not the major way.

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The major way is when oxygen
actually binds hemoglobin.

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In fact, we call that HbO2.

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And the name of that
molecule is oxyhemoglobin.

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So this is how the
majority of the oxygen

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is going to get
delivered to the tissues.

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And on the other side,
coming back from the tissue

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00:00:57,730 --> 00:01:00,597
to the lungs, you've got
dissolved carbon dioxide.

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A little bit of carbon
dioxide actually, literally

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comes just right in the plasma.

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But that's not the majority of
how carbon dioxide gets back.

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The more effective ways of
getting carbon dioxide back,

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remember, we have this
protonated hemoglobin.

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And actually
remember, when I say

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there's a proton
on the hemoglobin,

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there's got to be some bicarb
floating around in the plasma.

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And the reason that works is
because when they get back

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to the lungs, the proton, that
bicarb, actually meet up again.

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And they form CO2 and water.

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And this happens because
there's an enzyme called

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carbonic anhydrase inside
of the red blood cells.

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So this is where the carbon
dioxide actually gets back.

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And of course,
there's a third way.

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Remember, there's
also some hemoglobin

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that actually binds
directly to carbon dioxide.

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And in the process, it forms
a little proton as well.

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And that proton can
go do this business.

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It can bind to a
hemoglobin as well.

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So there's a little
interplay there.

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But the important ones I want
you to really kind of focus in

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00:02:04,390 --> 00:02:08,560
on are the fact that
hemoglobin can bind to oxygen.

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And also on this
side, that hemoglobin

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actually can bind to protons.

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00:02:13,800 --> 00:02:15,380
Now, the fun part
about all this is

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that there's a
little competition,

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a little game going on here.

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Because you've got,
on the one side,

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you've got hemoglobin
binding oxygen.

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And let me draw it twice.

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And let's say this top one
interacts with a proton.

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Well, that protons going to want
to snatch away the hemoglobin.

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And so there's a little
competition for hemoglobin.

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And here, the oxygen gets
left out in the cold.

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And the carbon dioxide does
the same thing, we said.

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Now, we have little hemoglobin
bound to carbon dioxide.

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And it makes a proton
in the process.

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But again, it leave
oxygen out in the cold.

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So depending on whether
you have a lot of oxygen

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around, if that's the kind
of key thing going on,

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or whether you have a lot
of these kinds of products

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the proton or the
carbon dioxide.

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Depending on which one you
have more of floating around

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in the tissue in the
cell, will determine

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which way that reaction goes.

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So keeping this
concept in mind, then I

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could actually step
back and say, well,

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I think that oxygen is affected
by carbon dioxide and protons.

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I could say, well, these two,
carbon dioxide and protons,

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are actually
affecting, let's say,

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are affecting the, let's say,
the affinity or the willingness

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of hemoglobin to bind,
of hemoglobin for oxygen.

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That's one kind of
statement you could

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make by looking at that
kind of competition.

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And another person come
along and they say,

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well, I think oxygen
actually is affecting,

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depending on which one,
which perspective you take.

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You could say, oxygen is
affecting maybe the affinity

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00:03:57,450 --> 00:04:03,530
of hemoglobin for the
carbon dioxide and proton

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00:04:03,530 --> 00:04:09,700
of hemoglobin for
CO2 and protons.

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So you could say it
from either perspective.

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And what I want to point
out is that actually,

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in a sense, both
of these are true.

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And a lot of times we
think, well, maybe it's

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just saying the
same thing twice.

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But actually, these are
two separate effects.

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And they have two
separate names.

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So the first one, talking about
carbon dioxide and protons,

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their effect is called
the Bohr effect.

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00:04:32,840 --> 00:04:36,230
So you might see that
word or this description.

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This is the Bohr effect.

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00:04:38,020 --> 00:04:41,170
And the other one, looking at
it from the other prospective,

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00:04:41,170 --> 00:04:43,210
looking at it from
oxygen's perspective,

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00:04:43,210 --> 00:04:45,430
this would be the
Haldane effect.

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00:04:45,430 --> 00:04:48,480
That's just the name
of it, Haldane effect.

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00:04:48,480 --> 00:04:50,770
So what is the Bohr effect
and the Haldane effect?

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Other than simply saying
that the things compete

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for hemoglobin.

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Well, let me actually bring
up a little bit of the canvas.

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And let's see if I
can't diagram this out.

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Because sometimes I think a
little diagram would really

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00:05:01,770 --> 00:05:03,530
go a long way in
explaining these things.

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So let's see if I can do that.

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Let's use a little graph and see
if we can illustrate the Bohr

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effect on this graph.

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So this is the partial
pressure of oxygen,

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how much is dissolved
in the plasma.

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And this is oxygen
content, which is to say,

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how much total oxygen
is there in the blood.

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And this, of course,
takes into account

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mostly the amount of oxygen
that's bound to hemoglobin.

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So as I slowly increase the
partial pressure of oxygen,

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see how initially,
not too much is

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going to be binding
to the hemoglobin.

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But eventually as a few
of the molecules bind,

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you get cooperativity.

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And so then, slowly the
slope starts to rise.

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And it becomes more steep.

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And this is all because
of cooperativity.

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Oxygen likes to bind where other
oxygens have already bound.

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, And then it's
going to level off.

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And the leveling off
is because hemoglobin

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is starting to get saturated.

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So there aren't too many
extra spots available.

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So you need lots and lots of
oxygen dissolved in the plasma

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to be able to seek out and
find those extra remaining

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spots on hemoglobin.

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So let's say we
choose two spots.

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One spot, let's say,
is a high amount

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of oxygen dissolved
in the blood.

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And this, let's
say, is a low amount

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of oxygen dissolved
in the blood.

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I'm just kind of choosing
them arbitrarily.

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And don't worry about the units.

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And if you were to think
of where in the body

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would be a high
location, that could

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be something like
the lungs where

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you have a lot of oxygen
dissolved in blood.

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And low would be, let's say,
the thigh muscle where there's

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a lot of CO2 but not so much
oxygen dissolved in the blood.

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So this could be two
parts of our body.

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And you can see that.

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Now, if I want to figure
out, looking at this curve

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how much oxygen is being
delivered to the thigh,

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then that's actually
pretty easy.

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I could just say, well, how much
oxygen was there in the lungs,

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or in the blood vessels
that are leaving the lungs.

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And there's this much
oxygen in the blood

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vessels leaving the lungs.

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And there's this much
oxygen in the blood

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vessels leaving the thigh.

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So the difference, whenever
oxygen is between these two

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points, that's the amount of
oxygen that got delivered.

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So if you want to figure out
how much oxygen got delivered

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to any tissue you can simply
subtract these two values.

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So that's the oxygen delivery.

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But looking at this, you
can see an interesting point

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which is that if you wanted to
increase the oxygen delivery.

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Let's say, you wanted
for some reason

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to increase it, become more
efficient, then really,

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the only way to
do that is to have

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the thigh become more hypoxic.

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As you move to the
left on here, that's

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really becoming hypoxic,
or having less oxygen.

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So if you become more
hypoxic, then, yes, you'll

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have maybe a lower point
here, maybe a point like this.

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And that would mean a
larger oxygen delivery.

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But that's not ideal.

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You don't want your
thighs to become hypoxic.

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That could start
aching and hurting.

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So is there another way to
have a large oxygen delivery

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without having any
hypoxic tissue,

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or tissue that has a low
amount of oxygen in it.

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And this is where the Bohr
effect comes into play.

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So remember, the
Bohr effect said

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that, CO2 and protons
affect the hemoglobin's

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affinity for oxygen.

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So let's think of a situation.

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I'll do it in green.

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And in this situation, where
you have a lot of carbon dioxide

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and protons, the
Bohr effect tells us

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that it's going to be harder
for oxygen to bind hemoglobin.

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So if I was to sketch
out another curve,

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initially, it's going to
be even less impressive,

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with less oxygen
bound to hemoglobin.

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And eventually, once the
concentration of oxygen

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rises enough, it will
start going up, up, up.

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00:08:53,070 --> 00:08:55,040
And it does bind
hemoglobin eventually.

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00:08:55,040 --> 00:08:57,000
So it's not like it'll
never bind hemoglobin

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00:08:57,000 --> 00:08:59,710
in the presence of carbon
dioxide and protons.

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But it takes longer.

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00:09:01,770 --> 00:09:05,355
And so the entire curve
looks shifted over.

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00:09:05,355 --> 00:09:09,340
These conditions of high
CO2 and high protons,

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00:09:09,340 --> 00:09:11,620
that's not really
relevant to the lungs.

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00:09:11,620 --> 00:09:14,540
The lungs are thinking,
well, for us, who cares.

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00:09:14,540 --> 00:09:16,450
We don't really have
these conditions.

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00:09:16,450 --> 00:09:18,890
But for the thigh,
it is relevant

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00:09:18,890 --> 00:09:20,960
because the thigh
has a lot of CO2.

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00:09:20,960 --> 00:09:23,050
And the thigh has
a lot of protons.

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00:09:23,050 --> 00:09:26,170
Again, remember, high
protons means low pH.

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00:09:26,170 --> 00:09:29,420
So you can think
of it either way.

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00:09:29,420 --> 00:09:33,190
So in the thigh, you're going
to get, then, a different point.

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00:09:33,190 --> 00:09:36,720
It's going to be on the green
curve not the blue curve.

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00:09:36,720 --> 00:09:40,310
So we can draw it at
the same O2 level,

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00:09:40,310 --> 00:09:42,230
actually being down here.

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00:09:42,230 --> 00:09:45,750
So what is the O2 content in the
blood that's leaving the thigh?

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00:09:45,750 --> 00:09:47,960
Well, then to do it
properly, I would say, well,

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00:09:47,960 --> 00:09:50,530
it would actually be over here.

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00:09:50,530 --> 00:09:52,420
This is the actual amount.

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00:09:52,420 --> 00:09:56,290
And so O2 deliver is actually
much more impressive.

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00:09:56,290 --> 00:09:57,360
Look at that.

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00:09:57,360 --> 00:10:01,127
So O2 delivery is increased
because of the Bohr effect.

216
00:10:01,127 --> 00:10:03,460
And if you want to know exactly
how much it's increased,

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00:10:03,460 --> 00:10:04,410
I could even show you.

218
00:10:04,410 --> 00:10:08,830
I could say, well, this
amount from here down to here.

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00:10:08,830 --> 00:10:11,820
Literally the vertical
distance between the green

220
00:10:11,820 --> 00:10:13,040
and the blue lines.

221
00:10:13,040 --> 00:10:17,650
So this is the extra oxygen
delivered because of the Bohr

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00:10:17,650 --> 00:10:19,050
effect.

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00:10:19,050 --> 00:10:22,670
So this is how the Bohr effect
is so important at actually

224
00:10:22,670 --> 00:10:25,710
helping us deliver
oxygen to our tissues.

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00:10:25,710 --> 00:10:28,710
So let's do the same thing,
now, but for the Haldane effect.

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00:10:28,710 --> 00:10:31,120
And to do this, we actually
have to switch things around.

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00:10:31,120 --> 00:10:34,050
So our units and our axes
are going to be different.

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00:10:34,050 --> 00:10:37,770
So we're going to have the
amount of carbon dioxide there.

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00:10:37,770 --> 00:10:41,490
And here, we'll do carbon
dioxide content in the blood.

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00:10:41,490 --> 00:10:43,880
So let's think through
this carefully.

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00:10:43,880 --> 00:10:45,930
Let's first start
out with increasing

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00:10:45,930 --> 00:10:48,670
the amount of carbon
dioxide slowly but surely.

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00:10:48,670 --> 00:10:50,830
And see how the content goes up.

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00:10:50,830 --> 00:10:53,600
And here, as you increase
the amount of carbon dioxide,

235
00:10:53,600 --> 00:10:56,529
the content is kind of
goes up as a straight line.

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00:10:56,529 --> 00:10:58,070
And the reason it
doesn't take that S

237
00:10:58,070 --> 00:10:59,960
shape that we had
with the oxygen

238
00:10:59,960 --> 00:11:03,220
is that there's no cooperativity
in binding the hemoglobin.

239
00:11:03,220 --> 00:11:05,710
It just goes up straight.

240
00:11:05,710 --> 00:11:07,400
So that's easy enough.

241
00:11:07,400 --> 00:11:10,010
Now, let's take two
points like we did before.

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00:11:10,010 --> 00:11:11,840
Let's take a point,
let's say up here.

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00:11:11,840 --> 00:11:15,040
This will be a high amount
of CO2 in the blood.

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00:11:15,040 --> 00:11:18,220
And this will be a low
amount of CO2 in the blood.

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00:11:18,220 --> 00:11:20,540
So you'd have a low amount,
let's say right here,

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in what part of the tissue?

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00:11:22,630 --> 00:11:25,780
Well, low CO2, that
sounds like the lungs

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00:11:25,780 --> 00:11:28,500
because there's not
too much CO2 there.

249
00:11:28,500 --> 00:11:31,750
But high CO2, it
probably is the thighs

250
00:11:31,750 --> 00:11:35,080
because the thighs like
little CO2 factories.

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00:11:35,080 --> 00:11:38,330
So the thigh has a high
amount and the lungs

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00:11:38,330 --> 00:11:39,080
have a low amount.

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00:11:39,080 --> 00:11:42,850
So if I want to look at the
amount of CO2 delivered,

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00:11:42,850 --> 00:11:43,850
we'd do it the same way.

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00:11:43,850 --> 00:11:47,340
We say, OK, well, the
thighs had a high amount.

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00:11:47,340 --> 00:11:50,990
And this is the amount of
CO2 in the blood, remember.

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00:11:50,990 --> 00:11:53,120
And this is the amount
of CO2 in the blood when

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00:11:53,120 --> 00:11:54,780
it gets to the lungs.

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00:11:54,780 --> 00:11:59,270
So the amount of CO2 that
was delivered from the thigh

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00:11:59,270 --> 00:12:01,860
to the lungs is the difference.

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00:12:01,860 --> 00:12:04,250
And so this is how
much CO2 delivery

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00:12:04,250 --> 00:12:05,670
we're actually getting.

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00:12:05,670 --> 00:12:10,560
So just like we had O2 delivery,
we have this much CO2 delivery.

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00:12:10,560 --> 00:12:12,640
Now, read over the
Haldane effect.

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00:12:12,640 --> 00:12:16,390
And let's see if we can actually
sketch out another line.

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In the presence of high
oxygen, what's going to happen?

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00:12:20,670 --> 00:12:23,270
Well, if there's a
lot of oxygen around,

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00:12:23,270 --> 00:12:26,650
then it's going to change
the affinity of hemoglobin

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00:12:26,650 --> 00:12:28,960
for carbon dioxide and protons.

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00:12:28,960 --> 00:12:34,500
So it's going to allow less
binding of protons and carbon

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00:12:34,500 --> 00:12:36,809
dioxide directly
to the hemoglobin.

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00:12:36,809 --> 00:12:38,850
And that means that you're
going to have less CO2

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00:12:38,850 --> 00:12:44,410
content for any given amount
of dissolved CO2 in the blood.

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00:12:44,410 --> 00:12:47,180
So the line still is a straight
line, but it's actually,

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00:12:47,180 --> 00:12:50,240
you notice, it's kind
of slope downwards.

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00:12:50,240 --> 00:12:52,820
So where is this relevant?

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00:12:52,820 --> 00:12:54,250
Where do you have
a lot of oxygen?

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00:12:54,250 --> 00:12:56,900
Well, it's not really
relevant for the thighs

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00:12:56,900 --> 00:12:59,370
because the thighs don't
have a lot of oxygen.

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00:12:59,370 --> 00:13:01,990
But it is relevant
for the lungs.

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00:13:01,990 --> 00:13:03,950
It is very relevant there.

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00:13:03,950 --> 00:13:07,640
So now you can actually say,
well, let's see what happens.

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00:13:07,640 --> 00:13:10,970
Now that you have high
O2, how much CO2 delivery

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00:13:10,970 --> 00:13:12,140
are you getting?

285
00:13:12,140 --> 00:13:14,160
And you can already see it.

286
00:13:14,160 --> 00:13:17,260
It's going to be more because
now you've got this much.

287
00:13:17,260 --> 00:13:21,290
You've got going all
the way over here.

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00:13:21,290 --> 00:13:23,650
So this is the new
amount of CO2 delivery.

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00:13:23,650 --> 00:13:24,770
And it's gone up.

290
00:13:24,770 --> 00:13:27,400
And in fact, you can even
show exactly how much

291
00:13:27,400 --> 00:13:30,010
it's gone up by, by simply
taking this difference.

292
00:13:30,010 --> 00:13:33,470
So this difference right
here between the two,

293
00:13:33,470 --> 00:13:36,280
this is the Haldane effect.

294
00:13:36,280 --> 00:13:38,900
This is the visual way
that you can actually

295
00:13:38,900 --> 00:13:40,880
see that Haldane effect.

296
00:13:40,880 --> 00:13:42,880
So the Bohr effect and
the Haldane effect, these

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00:13:42,880 --> 00:13:45,940
are two important
strategies our body

298
00:13:45,940 --> 00:13:49,720
has for increasing the
amount of O2 delivery and CO2

299
00:13:49,720 --> 00:13:51,770
delivery going back and
forth between the lungs

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00:13:51,770 --> 00:13:53,460
and the tissues.