Chapter 10 PHOTOSYNTHESIS

The sun is the energy source that keeps most organisms alive.
Photosynthesis converts radiant (solar) energy to chemical energy.
Almost all organisms depend on photosynthesis directly or indirectly.
Exception to this are the hot vent ecosystems ound at the !ottom o the oceans.
Producers are organisms that can make organic molecules rom inorganic su!stance.
They are also kno"n as autotrophs.
#ost producers are photoautotrophs and use light as energy source or manuacturing organic
compounds rom C$% and &%$.
$rganisms that cannot make their o"n organic compounds and must o!tain them rom other
organisms are called heterotrophs.
Consumers and decomposers are heterotrophs.
Photosynthesis is the !iochemical process that utili'es radiant energy rom sunlight to
synthesi'e car!ohydrates rom C$% and &%$ in the presence o chlorophyll.
CHLOROPLASTS
(n eukaryotes) photosynthesis occurs in chloroplasts) "hich are located mostly in the mesophyll
o the lea.
The mesophyll is the tissue in the middle o the lea in a cross section* it is ound in !et"een the
upper and lo"er epidermis.
#eso + middle
Phyll + lea
A typical cell has ,0 to -0 chloroplasts.
Chloroplasts are organelles !ound !y a dou!le mem!rane.
.imensions %/- 0m !y -/1 0m
The inner mem!rane encloses a luid/illed space called the stroma in "hich mem!ranous
lattened sacs) the thylakoids are stacked.
The thylakoid mem!rane encloses the thylakoid interior space.
The stacks o thylakoids are called grana (singular granum).
2sually there are -0 to 30 grana linked together !y arms in each chloroplast.
Thylakoids are part o an overlapping and continuous system o mem!ranes suspended in the
stroma.
4uilt in thylakoid mem!rane are the chlorophyll molecules and other pigments that capture light
energy.
The chlorophyll molecule is em!edded in the thylakoid mem!rane !y a long hydropho!ic
hydrocar!on tail
THE PATHAY O! PHOTOSYNTHESIS" A PRE#IE
Photosynthesis is the conversion o light energy into chemical !ond energy.

3C$% 5 1% &%$ C3&1%$3 5 3$% 5 3 &%$
Photosynthesis consists o t"o processes each "ith multiple steps. These processes are kno"n
as the light rea$tions and the Cal%in $y$le.
(n the light reactions) solar energy is converted to chemical energy.
The chloroplasts split "ater into hydrogen and oxygen.
The $% rom "ater is released to the atmosphere and restores the oxygen consumed
during respiration.
6ight provides the energy to transer electrons and hydrogen rom "ater to the acceptor
7A.P
5
.
7A.P
5
is reduced to 7A.P&.
ATP is also generated during the light reaction !y a process called
photophosphorylation.
ATP and 7A.P& are the products o the light/dependent reactions.
The oxygen in C$% is incorporated into car!ohydrates and "ater.
The car!on is reduced into car!ohydrates !y the addition o hydrogen atoms (electrons).
This process is called $ar&on 'i(ation.
THE NAT)RE O! S)NLI*HT
6ight is a orm o energy called electromagnetic energy or electromagnetic radiation.
6ight is composed o particles o energy that travel as "aves.
6ight is part o the ele$tromagneti$ spe$trum) the entire range o electromagnetic radiation.
8avelength is the distance rom one "ave peak to the next.
9isi!le light consists o a mixture o "avelengths ranging rom a!out ,:0 nm to 130 nm.
9iolet has the shortest "avelength and red the longest.
6ight also !ehaves as particles or packets o energy called photons.
The energy o the photon is inversely proportional to its "avelength; the shorter the "avelength
the greater the energy o the photon.
The atmosphere ilters out a large amount o radiation) !ut allo"s visi!le light to pass through.
PHOTOSYNTHETIC PI*+ENTS" THE LI*HT RECEPTORS
The spectrophotometer is used to measure the amount o light a su!stance has a!sor!ed.
The a&sorption spe$trum is a plot o the a!sorption o light o dierent "avelengths.
The a$tion spe$trum o photosynthesis sho"s ho" eective various "avelengths o light are in
carrying photosynthetic activity
STR)CT)RE O! THE CHLOROPHYLL +OLEC)LE,
There are several kinds o chlorophyll all o them containing a magnesium atom.
The molecule has t"o main parts; one captures the energy and the other holds the molecule in
place.
A complex ring called the porphyrin ring) made o <oined smaller rings made o car!on and
nitrogen a!sor!s light) and a hydrocar!on or lipid tail.
The porphyrin ring is very similar to the heme portion o the red pigment hemoglo!in in
red !lood cells.
The porphyrin ring o chlorophyll contains an atom o magnesium in its center.
The heme ring o hemoglo!in contains iron in its center.
The tail extends into the thylakoid mem!rane and is hydropho!ic.
The most important is chlorophyll a.
Chlorophyll ! is an accessory pigment that also participates in photosynthesis.
Chlorophyll a has a methyl group) C&,) on the porphyrin ring.
Chlorophyll ! has car!onyl group) C&$) on the porphyrin ring.
Chlorophyll a!sor!s light mostly in the !lue and red areas o the visi!le spectrum.
=reen light is not apprecia!ly a!sor!ed !ut relected !y chlorophyll.
Chloroplasts also have accessory pigments called $arotenoids) "hich are yello" and orange.
Carotenoids a!sor! light at ranges dierent rom chlorophyll and !roaden the spectrum o light
that provides energy or photosynthesis.
>ome carotenoids act as protectors that dissipate excess light that might damage chlorophyll.
To vie" the structural ormula o the chlorophyll molecules visit;
http;??users.rcn.com?<kim!all.ma.ultranet?4iologyPages?C?Chlorophyll.html
E-CITATION O! CHLOROPHYLL .Y LI*HT" THE LI*HT REACTIONS,
8hen a molecule a!sor!s one photon o light) one o its electrons is energi'ed and moves into
a higher energy level (excited state).
1. The excited electron may return to its original lo"er energy level (ground state) emitting
a less energetic photon. This is called luorescence. They may also release energy in
the orm o heat.
%. An electron acceptor molecule may accept the excited electron.
The only photons a!sor!ed are those "hose energy is e@ual to the dierence !et"een the
ground state and excited state.
A particular compound a!sor!s photons corresponding to speciic "avelengths) "hich is "hy
each pigment has a uni@ue a!sorption spectrum.
The energy o the a!sor!ed photon is converted to the energy o an electron raised rom the
ground level to an excited state.
Photosystems
Photosystems are light harvesting complexes made pigments and proteins. The various
pigments ena!le the photosystem to harvest light over a larger surace and a larger portion o
the spectrum.
Each photosystem consists o;
1. 6ight harvesting complexes made o chlorophyll) carotenoids) proteins and other
molecules.
%. A reaction center made o t"o special chlorophyll a molecules) P100 and P3:0 and a
primary electron acceptor.
T"o types o photosynthetic units present in chloroplasts make up photosystems kno"n as
photosystem ( and photosystem (() in order o their discovery.
Photosystem ( is a trimer; made o three su!units.
Each photosynthetic unit o photosystem ( consists o highly ordered groups o ...
%00 to ,00 molecules o chlorophyll a)
small amounts o chlorophyll !)
carotenoids "ith proteins attached)
a special rea$tion/$enter made o t"o molecules o chlorophyll a called P100.
primary electron acceptor molecules.
These photosynthetic units are also called antenna complexes.
The remaining photosystem pigment molecules are called antenna pigments.
The photosynthetic units o photosystem (( consists o...
%00 or more chlorophyll a)
/carotene attached to a protein)
a little chlorophyll !)
% molecules o chlorophyll o P3:0 in a reaction/center.
primary electron acceptor
The num!er 100 and 3:0 reer to the peak a!sorption "ith "avelengths o 100 and 3:0 nm.
4oth P100 and P3:0 are identical chlorophyll a molecules. Their slight dierence in a!sorption is
due to the associated proteins that aects the electron distri!ution in the molecules.
Aor a detail description o the structure o the t"o photosystems visit;
http;??users.rcn.com?<kim!all.ma.ultranet?4iologyPages?6?6ightBeactions.htmlCantenna
Light har%esting
8hen any antenna complex molecule traps a photon) it transers its energy rom pigment
molecule to pigment molecule until it reaches a particular molecule o chlorophyll a located in
the region o the complex called the reaction center.
The reaction/center molecule is the only one that can actually use the light energy.
A!sorption o a photon causes a transition o the chlorophyll molecule rom its ground state to
its excited state.
At the reaction center) a speciali'ed molecule called the primary ele$tron a$$eptor captures
an excited electron rom the reaction/center chlorophyll a molecule.
A redox reaction
The primary electron acceptor traps the energi'ed electron o chlorophyll a !eore it returns to
its ground state in the chlorophyll a molecule.
6ight energy is converted to chemical energy in the reaction center !y a series o electron
transer reactions.
An isolated chlorophyll molecule "ill lose its energy as heat and luorescent light as
the electron drops do"n to its original ground state.
.uring the light reactions o photosynthesis there are t"o possi!le routes or the electron lo";
cyclic and non/cyclic.
Non/$y$li$ ele$tron 'lo0
1. #any photons strike the photosystem molecules at the same time) "ith the energy o
excited electrons !eing transerred to"ard the P3:0 reaction center.
The energy !oosts an electron to a higher energy level.
The excited electron is taken !y the primary electron acceptor and P3:0 !ecomes oxidi'ed.
The oxidi'ed $hlorophyll a no" is a very strong reducing agent. (t has room or one
electron.
%. The electrons lost !y the P3:0 are replaced "ith electrons extracted rom "ater !y a protein
complex (en'yme) called the oxygen/evolving complex.
This process o splitting "ater to o!tain electrons is called photolysis.
The electrons extracted rom "ater are passed to tyrosine also kno"n as D.
D contains manganese) "hich is re@uired to split "ater.
T"o "ater molecules produce oxygen) $%) and our protons) &
5
.
,. The energi'ed electron is transerred to an electron acceptor) pheophytin) and then to P@)
plasto@uinone "ithin the thylakoid mem!rane.
P@ unloads the electrons to the iron/containing cytochromes ound in the thylakoid
mem!rane acing the stroma o the chloroplast.
Cytochromes pass the electron to plastocyanin) Pc) a copper containing protein.
These electrons are eventually donated to P100 in photosystem (.
Photosynthesis electron transport chain; Pheophytin E P@ E t"o cytochrome complexes E
Pc
-. 8hen electrons move along the transport system) protons move rom the stroma across the
thylakoid mem!rane to the interior thylakoid space !y chemiosmosis and ATP is
synthesi'ed.
This process is called non/cyclic photophosphorylation.
A proton gradient is created !et"een the stroma and the interior thylakoid space. The
greater proton concentration is in the interior thylakoid space.
8hen chloroplasts are illuminated) the p& in the stroma increases to :) and the p& in the
thylakoid space decreases to F) a dierence o three p& units.
The interior thylakoid space has a!out 1000 old higher concentration o &
5
.
F. The lo" energy electron no" reaches the end o the chain and is donated to P100) the
chlorophyll a molecule in the reaction center o photosystem (. The chlorophyll a molecule
!ecomes reduced.
Pigment molecules in the antenna complex o photosystem ( a!sor! photons and pass their
energy to the reaction center) P100) and an electron is energi'ed (excited).
3. The excited electrons are passed to an acceptor molecule) P-,0) "hich transers it to three
iron/sulur/containing proteins and then to errodoxin) Ad) a mem!rane/!ound iron/
containing protein.
Aerrodoxin transers the electron to AA. reducing it to AA.&%.
AA.&% then reduces 7A.P
5
to 7A.P& "ith the help o the en'yme 7A.P reductase. T"o
electrons are re@uired to reduce 7A.P
5
to 7A.P&.
7A.P& is released into the stroma o the chloroplast.
Electrons rom photosystem (( replace the missing electrons o P100.
>econd electron transport chain; Ad E AA.&% E 7A.P
5
reductase E 7A.P&
The energy stored in 7A.P& "ill provide reducing po"er or the synthesis o car!ohydrates.
Cy$li$ ele$tron 'lo0
2nder certain conditions) the excite electrons lo" !ack rom errodoxin to the cytochrome
complex and rom there continues on to P100.
This lo" generates ATP again.
The Calvin cycle is the next part o photosynthesis and re@uires more ATP than 7A.P&. The
cyclic lo" makes up the dierence.
Aollo" the electrons* a summary;
e
/
removed rom "ater (photolysis) are passed on to photosystem (() then to P@) the cytochrome
chain) photosystem () P-,0) errodoxin and inally 7A.P&.
(n the process photophosphorylation occurred through chemiosmosis.
This entire process is also kno"s as non/cyclic photophosphorylation) a continuous
linear process.
An alternate path o electrons is called cyclic photophosphorylation) "hich produces
ATP !ut no 7A.P&.
The electrons in can !e transerred to the electron transport chain !et"een the t"o
photosystems and !ack into photosystem (.
Comparison &et0een $hloroplasts and mito$hondria,
Similarities"
Chloroplasts and mitochondria generate ATP !y the same mechanism o chemiosmosis.
An electron transport chain pumps electron across a mem!rane and a proton gradient is
created.
Potential energy is stored in the orm o &
5
gradient.
4oth organelles have !uilt in their mem!ranes a similar ATP/synthase complex that couples the
lo" o electrons do"n the gradient "ith the phosphorylation o A.P.
Electron carriers) cytochromes and ATP synthase are very similar in !oth organelles.
1i''eren$es"
#itochondria transer chemical energy rom ood to ATP* chloroplasts convert radiant energy to
chemical energy.
(n mitochondria) the high/energy electrons are extracted rom ood molecules. Aood
molecules are oxidi'ed.
Chloroplasts capture light energy) "hich is used to excite lo"/energy electrons extracted
rom "ater and drive them to the top o the chain.
The spatial organi'ation o chemiosmosis is dierent in !oth organelles.
(n the mitochondrion) the electrons are pumped out o the matrix into the intermem!rane
space. ATP is synthesi'ed in the mitochondrial matrix.
(n the chloroplast) the electrons are pumped into the thylakoid space rom the stroma.
ATP is synthesi'ed in the stroma.
THE CAL#IN CYCLE 2LI*HT IN1EPEN1ENT REACTIONS OR 1AR3 REACTIONS4
The Calvin cycle is a cyclic meta!olic path"ay.
Car!on enters the cycle in the orm o car!on dioxide and leaves in the orm o car!ohydrate.
ATP is the source o energy and 7A.P& is the reducing po"er !y adding high/energy electrons
to the car!ohydrate.
The car!ohydrate produced directly rom the Calvin cycle is *5P or gly$eraldehyde/5/
phosphate.
Car!on ixation reers to the incorporation o C$% into an organic molecule.
.uring the car!on ixation reactions) ATP and 7A.P& are used to manuacture car!ohydrate
molecules rom C$%.
#ost plants use the Calvin cycle) also kno"n as the C, cycle) to ix car!on.
#ain steps o the Calvin cycle;
1. Car&on 'i(ation; Three molecules o C$% rom air com!ine "ith , molecules o
ri!ulose 1)F/!iphosphate) Bu4P) "ith the aid o the en'yme ru&is$o (Ru.P
$ar&o(ylase).
Bu4P is a F/C sugar constantly !eing ormed in the cycle.
%. The resulting 3/C unsta!le complexes are immediately split into 3 molecules o 5/
phosphogly$erate 25P*A4) the irst sta!le compound ormed in photosynthesis.
,. Redu$tion; Each molecule o the 3 molecules o ,P=A receives a phosphate rom ATP
and !ecome 1), !iphosphoglycerate.
-. 7A.P& supplies electrons to reduce the car!oxyl group o 1), !iphosphoglycerate to the
car!onyl group o glyceraldehyde , phosphate) =,P. =,P is the same sugar ormed in
glycolysis !y the splitting o glucose.
F. The 7A.P& and ATP supply the energy to convert the ,P=A to 3 molecules o
glyceraldehyde ,/phosphate (=,P)) a ,/C sugar phosphate. An inorganic phosphate
leaves the !iphosphoglycerate molecules in the process.
3. $ne o the =,P exits the cycle to !e used !y the plant cell. The other ive are must !e
recycle to regenerate the three molecules o Bu4P
1. Regeneration o' Ru.P6 the CO7 a$$eptor; The other ive ,/C molecules o
glyceraldehyde ,/phosphate (=,P) are restructured in a series o complex reactions and
!ecome three F/C molecules o Bu4P) the sugar used in the irst step o the Calvin
cycle. Three ATP molecules are used in this process o regeneration o Bu4P.
:. This leaves a net gain o one glyceraldehyde ,/phosphate) =,P) "hich can contri!ute to
either an increase in the car!ohydrate content o the plant (glucose) starch) cellulose)
etc.) or can !e used in path"ays that lead to the net gain o lipids and amino acids.
G. Aor the synthesis o one =,P) the Calvin cycle consumes a total o nine molecules o
ATP ad six molecules 7A.P&. The light reaction generates these molecules.
S)++ARY
6ight/dependent reaction
87 H7O 9 87 NA1P
9
9 8: A1P 9 8: Pi ; O7 9 87 NA1PH 9 8: ATP 9 87 H
9
6ight/independent reaction
87 NA1PH 9 87 H
9
9 8: ATP 9 ; CO7 C;H87O; 9 87 NA1P
9
9 8: A1P 9 8: Pi 9 ; H7O
4y canceling the common items on the a!ove reactions "e o!tain the overall reaction o
photosynthesis.
; CO7 9 87 H7O C;H87O; 9 ; O7 9 ; H7O
ALTERNATI#E +ECHANIS+S O! CAR.ON !I-ATION
.ehydration is a danger or terrestrial plants.
Plants have to compromise !et"een photosynthesis and transpiration.
$pening o the stomata allo"s C$% to enter the mesophyll o' the lea') the site o
photosynthesis.
$pening o the stomata allo"s the escape o "ater rom the lea; transpiration.
The pro&lem
(n hot) dry days) most plants close their stomata in order to conserve "ater.
The concentration o C$% in the spaces in the mesophyll o the lea !egins to decrease as soon
as the stomata close) and the concentration o $% !egins to increase.
1) Photorespiration
The irst organic product o car!on ixation is a ,/C compound called ,/phosphoglycerate.
Plants that produce this ,/C compound are called C5 plants.
#any plants do not yield as much car!ohydrate as expected) especially during hot summer
spells.
1. $n hot) dry days plants close their stomata to conserve "ater.
%. $nce the stomata close) the plant uses rapidly the availa!le C$% and produces $%)
"hich accumulates in the chloroplasts.
,. Bu!isco is the en'yme responsi!le or !inding C$% to Bu4P to make P=A
(phosphoglyceraldehyde).
-. $% competes "ith C$% or the active !inding site o Bu!isco.
F. 8hen $% !inds to Bu!isco some intermediates o the Calvin cycle are degraded to C$%.
Phosphoglycolate) a %/C is produced "ith the consumption o a lot o energy.
3. Phosphoglycolate 5 $% 5 ATP E P=A 5 C$% 5 A.P 5 %Pi
1. Photorespiration occurs in chloroplasts) peroxisomes) and mitochondria o plant cells.
This process is called photorespiration !ecause* it occurs in the presence o light) re@uires $%
and produces C$% and &%$.
7o ATP is produced during photorespiration and the removal o Calvin cycle intermediates
reduces yield. 7o car!ohydrates) ood) are produced.
Bi!ulose/1)F/bis/phosphate car!oxylase?oxygenase
Photorespiration may !e an evolutionary relic rom a time "hen the concentration o oxygen in
the atmosphere "as very lo" and ru!isco) in its evolution) did not develop a "ay o excluding $%
rom its active site.
Bu!isco may !e conserving this original ainity or $%) even though the concentration o $% in
the atmosphere is no" very high) 1:H.
Bice) "heat and soy!ean may lose up to F0H o its yield in photorespiration.
Aor a good explanation o the photorespiration chemical reaction see;
http;??""".steve.g!.com?science?photorespiration.html
C< plants
(t is also called the Hat$h/Sla$k path0ay.
C$% is not very a!undant in the atmosphere; =,=5> o atmosphere.
.iusion o gases occurs only across moist suraces
6eaves and other structures are covered "ith "aterproo su!stances to prevent dehydration.
Entry o C$% is limited to the stomata) "hich lead to the interior o the lea) the mesophyll.
#esophyll cells contain chloroplasts and carry on photosynthesis.
8hen conditions are hot and dry) the stomata close to prevent excessive "ater loss) also
preventing C$% rom entering the lea interior.
C- plants ix car!on in the orm o a 'our/$ar&on $ompound !eore entering the Calvin
cycle.
C- plants have a uni@ue lea anatomy; &undle sheath $ells and mesophyll $ells.
The !undle sheath cells orm a tight ring around the veins and veinlets o the lea.
The stomata are oten closed in hot and dry climates "hen the maximum amount o light is
availa!le or photosynthesis.
1. C- reactions take place in the mesophyll o the cell.
%. The C$% is ixed into the -/C o(aloa$etate !y the en'yme PEP car!oxylase) "hich is
converted to malate and re@uires 7A.P&.
,. #alate moves into the chloroplasts o the &undle sheath $ell)
-. C$% is removed rom malate and pyru%ate is produced.
F. The generated C$% then enters the Calvin cycle.
3. Pyruvate returns to the mesophyll "here is converted to phosphoenolpyru%ate6 PEP.
The key component o the C- path"ay is the en'yme PEP $ar&o(ylase) an en'yme ound in
the mesophyll cells o leaves.
Compared to ru!isco) PEP $ar&o(ylase has a %ery high a''inity or C$% even at very lo"
concentrations.
C- must have the stomata partially open in order to conserve "ater. This condition causes the
concentration o C$% decrease and that o $% increase.
PEP car!oxylase cataly'es the reaction !y "hich C$% reacts "ith the ,/C compound
phosphoenolpyruvate to orm oxaloacetate) a -/C molecule.
8hen light is a!undant) the rate o photosynthesis is limited !y the concentration o C$%.
PEP car!oxylase can ix car!on "hen ru!isco cannot.
4ecause PEP car!oxylase has such a high ainity or C$%) C- plants tolerate higher
temperatures and higher light intensities) lose less "ater !y transpiration) and have higher rate
o photosynthesis and gro"th than plants that use only the Calvin cycle.

>everal thousand species o plants exhi!it the C- meta!olic path"ay e. g. sugar cane) corn and
cra!grass.
At lo"er temperatures and light intensities) C, plants are avored !ecause they do not re@uire as
much energy to ix C$%
C- and C, occur at dierent locations (cells) in the lea.
THE CRASS)LACEAN ACI1 +ETA.OLIS+ OR CA+,
CA# is similar to C- path"ay.
.esert plants use CA# in photosynthesis) e.g. succulents) cacti) euphor!s) orchids and
mem!ers o the pineapple amily.
The stomata $lose during the day to prevent excessive evaporation and open at night.
PEP car!oxylase ixes C$% at night in the mesophyll o the cell "hen the stomata are open.
$xaloacetate is ormed "hich is converted to malate and stored in cell vacuoles.
8hen light is availa!le) C$% is removed rom malate !y a decar!oxylation reaction) and is made
availa!le to the Calvin cycle.
CA# and C, occur at dierent times "ithin the same cell.
C, and C- plants sho" a spatial separation o meta!olic steps.
C, and CA# plants sho" a temporal separation o meta!olic steps.
C- and CA# plants sho" a temporal separation o meta!olic steps.
>ee; http;??""".steve.g!.com?science?photorespiration.html
$nly the green cells o the plant are autotrophic. All other cells consume !ut do not produce
ood) e. g. roots) trunk "ood and most parts o !ranches.
#ost plants manage to produce more organic material than they need or respiration and
gro"th.
The extra car!ohydrates are stored in roots) stems) ruits) seeds) and leaves.
$n the glo!al scale) the estimate ood produced !y plants is 130 !illion metric tons o
car!ohydrate per year.
1 metric ton + 1.1 tons or 1)000 kg or %)%00 pounds.
Photosynthesis provides the energy and !uilding material or ecosystems.