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Lesson 15: Krebs (citric acid) cycle and oxidative phosphorylation- Krebs (citric acid) cycle and oxidative phosphorylation questions
- Oxidative phosphorylation questions
- The citric acid cycle
- Krebs / citric acid cycle
- Regulation of pyruvate dehydrogenase
- Regulation of Krebs-TCA cycle
- Electron transport chain
- Oxidative Phosphorylation: The major energy provider of the cell
- Oxidative phosphorylation and chemiosmosis
- Regulation of oxidative phosphorylation
- Mitochondria, apoptosis, and oxidative stress
- Calculating ATP produced in cellular respiration
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Electron transport chain
Explore the electron transport chain's role in cellular respiration. Learn how NADH and FADH2 contribute to ATP production. Discover how energy is released when electrons transition from high to low energy states, creating a proton gradient that drives ATP synthase. Dive into the cutting-edge research in this field. Created by Sal Khan.
Want to join the conversation?
- Why do the H+ want to go back into the matrix? I know it must due to entropy but aren't the inside (matrix) and the outside part two closed systems?(15 votes)
- Diffusion causes them to want to go back across the membrane. They "want" to go down the concentration gradient.(10 votes)
- When protons are pumped from the matrix to the outer compartment, Sal explains they want to come back in (to steer away from the concentration gradient). What prevents them from escaping the gradient the other way and go back into the cytoplasm?(5 votes)
- What prevents hydrogen protons from escaping from the mitochondria and entering the cytoplasm is the lipid bi-layer membrane of the mitochondria. As you might now, these lipids are uncharged. They allow for easy passage of uncharged particles but not for charged ones. Normally charged particles enter into membranes through pores. ATP-synthase, is similar to such a pore. It allows the proton gradient to escape but only into the matrix(10 votes)
- Does this all happen in our body?(2 votes)
- yes this does happen in out body, in the mitochondria which is in the cytoplasm.(9 votes)
- AtSal talks about ATP synthase. I was wondering what is the difference between ATP synthase and ATPase or is there no difference? 9:57(3 votes)
- ATP synthase is an enzyme that uses the proton gradient to create ATP from ADP and inorganic phosphate.
An enzyme that is an ATPase is an enzyme that hydrolyzes ATP to do its job.(9 votes)
- How do the H+ ions know to pass through the ATP synthase? can they not diffuse out through the membrane? is it something to do with the charges on the protein channel? thanks :)(5 votes)
- We also know that for each electron that NADH and FADH2 deliver to the protein complex that belong to the electron transport chain, and amount H+ will be pump out to the inner membrane space. So that is how protons get to the inner membrane space and gradient forms.(1 vote)
- I think according to new research that each NADH produces 2.5 ATP and each FADH2 produces 1.5 ATP right?(4 votes)
- Yes, they also mention that and further discuss the ETC in this article, if you're interested: https://www.khanacademy.org/science/biology/cellular-respiration-and-fermentation/oxidative-phosphorylation/a/oxidative-phosphorylation-etc(2 votes)
- how do the electrons split from the NADHs to go through this process?(3 votes)
- The NADH gets oxidized by the protein complex called 'NADH dehydrogenase' (very 'original' name :P) into NAD+, and 2 H+ ions, and 2 e- (or electrons), so the balanced equation is: NADH -> NAD^+ + 2H^+ + 2e^-. btw... these electron are a small percentage of our 'free radicals', or extremely reactive electrons, that can cause major damage to cells, proteins, and DNA (cancer causing in other words)... but are also needed for the processes of cellular respiration :)(4 votes)
- what exactly is the matrix of the mitochondria?(3 votes)
- The mitochondrial matrix is enclosed within the inner mitochondrial membrane. It is really viscous compared to the cytoplasmic fluid. The mitochondrial circular DNA, some of the respiratory enzymes and the ribosomes are present in the matrix.(2 votes)
- why is it that the number of H20 molecules produced in aerobic cellular respiration, and the number of H20 molecules that theoretically should be produced according to the formula ( C6H12O6 + 6O2 --> 6H2O + 6CO2 + energy) do not match up? according to the formula, 6 molecules of water should be produced for every molecule of glucose. but in reality, 14 molecules are produced, and then 6 are used up during the krebs cycle which gives a net of 8 molecules of water produced. Why is this so?(3 votes)
- How long does the whole cellular respiration process take?(2 votes)
- A good question, but difficult to answer. This is because "how long" is a bit ill-defined. If you mean what is the time a particular carbon atom in glucose takes on appearing as carbon dioxide on entering the cell, the answer is variable. It can be seconds, milliseconds in high demand cases when there is a rapid conversion of glucose into successive metabolites. If the organism is at leisure, it is usually in seconds or minutes. If there is already a lot of reserve glucose in the cell as glycogen or otherwise, it may also be several minutes.(3 votes)
Video transcript
After being done with glycolysis
and the Krebs Cycle, we're left with 10
NADHs and 2 FADH2s. And I told you that these are
going to be used in the electron transport chain. And they're all sitting in the
matrix of our mitochondria. And I said they're going to be
used in the electron transport chain in order to actually
generate ATP. So that's what I'm going to
focus on in this video. The electron transport chain. And just so you know, a lot
of this stuff is known. But some of the details
are actually current areas of research. People have models and
they're trying to substantiate the models. But things are happening at such
a small scale here that people can just look at the
evidence, some of which is indirect, and say, this is
probably what's happening. Most of this is very well
established, but some of the exact mechanisms-- for example,
how exactly some of the proteins work-- aren't
completely understood. So I think it's very important
for you to understand that this is at the cutting edge,
that you're already there. So the basic idea here is that
the NADHs-- and that's where I'll focus. FADH2 is kind of
the same idea. Although its electrons
are just at slightly lower energy state. So they won't produce
quite as many ATPs. Each NADH is going to be-- as
you'll see-- indirectly responsible for the production
of three ATPs. And each FADH2, in a very
efficient cell, in both of these cases, will be indirectly
responsible for the production of two ATPs. And the reason why this guy
produces fewer ATPs is because the electrons that he has
going into the electron transport chain are at a
slightly lower energy level than the ones from NADH. So in general, I just
said indirectly. How does this whole
business work? Well in general, NADH, when it
gets oxidized-- remember, oxidation is the losing of
electrons or the losing of hydrogens that happen
to have electrons. We can write its half
reaction like this. Its oxidation reaction
like this. You'll have some NAD plus, which
you can then go and use back in the Krebs Cycle
and in glycolysis. You have some NAD plus, you'll
have a proton, a positive hydrogen ion is just a proton. And then you'll have
two electrons. This is the oxidation of NADH. It's losing these
two electrons. Oxidation is losing electrons. OIL RIG. Oxidation is losing electrons. Or you can imagine it's losing
hydrogens, from which it can hog electrons. Either one of those
is the case. Now this is really the
first step of the electron transport chain. These electrons are transported
out of the NADH. Now, the last step of the
electron transport chain is you have two electrons-- and you
could view it as the same two electrons if you like--
two electrons plus two hydrogen protons. And obviously if you just add
these two together, you're just going to have two hydrogen
atoms, which is just a proton and an electron. Plus one oxygen atom, so
I could say one half of molecular oxygen. That's the same thing as
saying one oxygen atom. And you're going to produce--
if I have one oxygen and two complete hydrogens, I'm
left with water. And you could view this, we're
adding electron or we're gaining electrons to oxygen. OIL RIG. Reduction is gaining
electrons. So this is the reduction
of oxygen to water. This is the oxidation
of NADH to NAD plus. Now, these electrons that are
popping out of-- these electrons right here-- that are
popping out of this NADH. And when they're in
NADH they're at a very high energy state. And what happens over the
course of the electron transport chain is that these
electrons get transported to a series of, I guess
you could call them transition molecules. But these transition molecules,
as the electrons go from one to the other,
they go into slightly lower energy states. And I won't even go into the
details of these molecules. One is coenzyme Q,
and cytochrome C. And then they eventually end
up right here and they are used to reduce your
oxygen into water. Now every time an electron
goes from a higher energy state to a lower energy state--
and that's what it's doing over the course of this
electron transport chain-- it's releasing energy. So energy is released when you
go from a higher state to a lower state. When these electrons were in
NADH, they were at a higher state than they are when they
bond to coenzyme Q. So they release energy. Then they go to cytochrome
C and release energy. Now that energy is used to
pump protons across the cristae across the inner
membrane of the mitochondria. And I know this is all very
complicated sounding. And this is the cutting edge. So it maybe should sound
a little complicated. Let me draw a mitochondria. So let me draw a small
mitochondria just so you know where we're operating. That's its outer membrane. And then its inner membrane,
or its cristae, would look like that. And let me zoom in
on the membrane. So let's say if I were to
zoom in right there. So if I were to zoom that out,
that box would look like this. You have your crista here. And I'm going to
draw it thick. So I'm zooming in. This green line right
here, I'm going to draw it really thick. I'm going to color it in with
the green, just like that. And then you have your
outer membrane. This outer membrane, I
can do it up here. And I'll just color it in. You don't even have to see the
outside of the outer membrane. Right here, this space right
here, this is the outer compartment. And then we learned in the last
video, this space right here is the matrix. This is where our Krebs
cycle occurred. And where a lot of our NADH,
or really all of our NADH, is sitting. So what happens is, every time
NADH gets oxidized to NAD plus, and the electrons keep
transferring from one molecule to another, it's occurring in
these big protein complexes. And I'm not going to go into
the details on this. So each of these protein
complexes span-- so let's say that's a protein complex where
this first oxidation reaction is occurring and releasing
energy. And then let's say there's
another protein complex here, where the second oxidation
reaction is occurring and releasing energy. And these proteins are able to
use that energy to essentially pump-- this might all seem
very complicated-- to essentially pump hydrogens
into the outer membrane. It actually pumps hydrogen
protons. And let me be very clear. Hydrogen protons into
the outer membrane. And every one of these reactions
pump out a certain number of hydrogen protons. So by the end of the electron
transport chain, or if we just followed one set of electrons,
by the time that they've gone from their high energy state in
NADH to their lower energy state in water, by the time
they've done that, they've supplied the energy to these
protein complexes that span our cristae to pump hydrogen
from the matrix into the outer membrane. So really the only byproduct of
the oxidation of NADH into, eventually, water, or the
oxidation of NADH and the reduction of oxygen into
water, isn't ATPs yet. It's just this gradient where we
have a lot higher hydrogen proton concentration in the
outer compartment than we do in the matrix. Or you could say that the outer
compartment becomes a lot more acidic. Remember acidity is just
hydrogen proton concentration, the concentration of
hydrogen protons. So the byproduct of all of this
energy is used to really just pump these protons into
the outer membrane. So you have two things. The outer membrane becomes
more acidic than the matrix inside. Maybe we could call
that basic. And obviously these are all
positively charged particles. So there's actually an electric
gradient, an electric potential between the
outer membrane and the inner membrane. This becomes slightly negative,
that becomes slightly positive. These guys wouldn't naturally
do this on their own. If this is already acidic and
it's already positive, left to its own devices, these more
protons wouldn't be entering. And the energy to do that is
supplied by electrons going from high energy state in NADH
to going to a lower energy state, eventually, on the
oxygen in the water. That's what's happening. But essentially all that's
happening is protons being pumped from the matrix into
the outer compartment. Now once that gradient
forms, these guys want to get back in. These guys want to get
back into the matrix. And that is where the
ATPs are formed. So there's a protein that
also spans this. Let me draw. Remember this is all this inner
membrane right here. Let me just draw it a little
bit bigger right here. So that's our inner membrane,
our cristae right there. There's a special protein
called-- and I'll show you actually a better diagram of
what looks like in a second-- called ATP synthase. And what happens is, remember
because of the electron transport chain, we have all
of these hydrogen ions up here, all of these
protons really. All they are is a proton. That really want to get back
into the matrix down here. But they can't. This crista is impermeable to
them so they have to find a special way to get through. They were able to go the reverse
direction through the special protein complexes. Now they're going to
go back into the matrix through ATP synthase. So they're going
to go back, but something interesting happens. And this is really an area of
current research where people think they know how it works
but they're not sure. Because you can't just take
these proteins apart and watch them operate like you can a
regular mechanical engine. These are ultra-small and they
have to be in a living system. And they have to have the
right conditions. And you can't even-- it's hard
to see hydrogen protons. These are ultra-small things
that are, pretty much, you can't see them. But what happens, the current
model is, as these enter, as these go through
my ATP synthase there's actually an axle. So you can kind of view
this as a housing. And then there's an axle. And this is all just
a big protein. And there's an axle and then
there's another part of the synthase down here. So you can imagine, this is
kind of mind-blowing. That something this fancy is
occurring on the membranes of pretty much all living
systems' cells. It's not just eukaryotes. Even prokaryotes do this. They don't do it in their
mitochondria; they do it in their cellular membrane. But it's a pretty neat thing. And what happens is, as these
go through, you can kind of imagine as water flowing
through a turbine. It mechanically causes
this structure in the middle to spin. To actually spin. This is the current thinking. And this thing is all uneven. It's not, like, this
nice tube. It'll look all crazy
like that. And what happens is that an ADP
molecule-- let's say that this is the A part of the ADP. And then you have two
phosphate groups. It'll attach to one part
of this protein. And maybe a phosphate will
just randomly attach to another part of this protein. Just like that. So right now it's just
ADP and a phosphate. But as this inner axle turns--
because it's not a symmetrical tube, it has different things
sticking out that have different amounts of atomic
charge and it's going to play with this outer housing
right here. And so as this turns, the outer
housing, because of just the proteins bumping against
each other and electrical charge and whatever else, it's
going to squeeze the ADP and the phosphate together to form,
actually form, ATP. And actually the current
thinking is that it does it on three different sites
simultaneously. So as this spins around, ADP and
phosphate groups kind of show up on the inside
of this housing. You could imagine
it like that. And I don't even know if
it's on the inside. But they show up
on the housing. And as this thing spins around,
it stretches and pulls on this outer part and pushes
these two things together. So it's using the energy from
this proton gradient to drive this axle. And because it's all strange, it
does all these distortions on this outer part and
actually pushes the two ATPs together. So when you start off with your
10 NADHs, it'll provided just enough energy and just
enough protons to put into the outer membrane that when they
go back through our ATP synthase-- you could almost
view it as an ATP synthase motor-- just based on people's
observations they see that this will produce, on
a per-NADH level, roughly three ATPs. On a per-FADH2 level,
roughly two ATPs. And I've said multiple times
in the videos, this is kind of an ideal. That a lot of times, maybe
you'll have some protons leak, so their energy can't be
captured properly. Or maybe some of these electrons
might somehow jump the gun or jump some steps, so
some of the energy gets lost. So you don't always have a
completely efficient system. And just so you believe that
this is actually occurring on our membrane, there's
actual visual depictions of these proteins. This is the actual protein
structure of ATP synthase right here. That is actually ATP synthase. And as you can see, there's
this piece right here that holds this part and that part. You can kind of imagine it
relatively stationary. The hydrogen comes
through here. The axle gets spun. And as the axle gets spun, ADP
and phosphate groups that are lodged inside this
F1 part of the protein, get pushed together. You have to put energy into the
reaction in order to make them stick together. But they get pushed together by
the protein itself as this axle turns around. And this axle turns around
from the energy of the hydrogen going. I don't even know what the
mechanics would look like. But you could imagine-- in my
head I imagine, the simplest thing is a windmill. Or not a windmill, as maybe some
type of water turbine or maybe the simplest thing
is, if you have something like that. I don't know if that's
what the protein actually looks like. If you have any kind of
thing passing by, it's going to spin it. It's going to spin
it like that. And you could be more creative
if you want to change the angle of the spin and whatnot. And that's all, people are
really still trying to understand this at a deeper
and deeper level. But for your purposes,
especially in an introductory biology level, you just have to
realize that two things are happening in the electron
transport chain. Electrons are moving from the
NADHs and the FADH2s to eventually show up and
reduce the oxygen. And as they do that, they're
releasing energy as they go from one molecule to another. They're going to lower
energy states. That energy is used to pump
hydrogen protons into the outer compartment of
the mitochondria. And then that gradient, those
hydrogen protons want to get back into the matrix of
the mitochondria. So as they go back in, that
drives this ATP synthase engine, which actually
produces the ATP. So just like we said in the
past, when you have 10 of these, on average-- let me say
this way-- on average each NADH is going to
produce 3 ATPs. Not directly. It produces enough of a gradient
of hydrogen protons to produce 3 ATPs in
the ATP synthase. And each FADH2, on average,
produces enough of a hydrogen gradient to produce 2 ATPs. So if we come in with 10 NADH,
they're going to produce-- in this ideal world-- 30 ATP. And then our 2 FADH2s are
going to produce 4 ATP. And then if you remember from
glycolysis, we had 2 net ATPs directly produced. And from the Krebs cycle we had
2 ATPs directly produced. So then you have 4 from
glycolysis and Krebs, and that gets us, once again, to our
magic 38 ATPs from one molecule of glucose. And now, I think you have a
pretty good grasp of cellular respiration.