FISIOLOGIA VEGETAL

FOTOSSÍNTESE

Profa Dra Ana Cardoso Clemente F. F. de Paula

DCA/ IFMG- Bambuí
The biosynthetic
pathway of chlorophyll.
The pathway begins with
glutamic acid, which is
converted to 5-
aminolevulinic acid
(ALA). Two molecules of
ALA are condensed to
form porphobilinogen
(PBG). Four PBG
molecules are linked to
form protoporphyrin IX.
The magnesium (Mg) is
then inserted, and the
light-dependent
cyclization of ring E, the
reduction of ring D, and
the attachment of the
phytol tail complete the
process.
 Carotenóides

Carotenos – α e β, licopeno
Xantofilas - luteína, neoxantina, violaxantina
Derivados dos carotenos
As xantofilas contém átomos de oxigénio e os carotenos apenas C e H
 Ficobilinas
Existem nas algas vermelhas e nas cianobactérias
Diferenças em relação à clorofila:
não possuem fitol
não contêm Mg
arranjo dos grupos tetrapirrólicos não é cíclico

Figure 19.32. Structure of a Phycobilisome. (A) Electron micrograph of

phycobilisomes from a cyanobacterium (Synechocystis). (B) Schematic
representation of a phycobilisome from the cyanobacterium Synechocystis 6701.
Rods containing phycoerythrin (PE) and phycocyanin (PC) emerge from a core made
of allophycocyanin (AP) and allophycocyanin B (APB). The core region binds to
 Movimento de folhas -
Movimento cloroplastos
• Fotossíntese: Um pouco de história

• van Helmont (1648) - aumento da massa do solo: H2O

• Hales (1727) - plantas extraem matéria do ar
• Priestley (1771) - as plantas libertam um tipo de ar que permite a
• Combustão
• Ingehousz (1779) - o poder “purificador”das plantas é devido à
• influência do sol nas suas partes verdes
• Senebier (1782) - CO2, ao qual chamou “ar fixo” é consumido
• durante a fotossíntese
• de Saussure (1804) - massas combinadas de matéria produzida
• pelas plantas e o O2 que elas libertam são maiores que a massa de
• CO2 que elas consumem
• Mayer (1842) - os organismos fotossintéticos convertem energia
• luminosa em energia livre química
 Fotossíntese
Fotoquímica Carboxilação
LUZ CO2

ATP
CICLO DE
HO
2 FOTOFOSFORILAÇÃO NADPH CALVIN
FOTÓLISE DA ÁGUA

O2 C6H12O6
 Formação de O2 a partir da água

Formação de O2 em membranas fotossintéticas

expostas a uma série de flashes de luz

Não há libertação de O2 nos dois primeiros flashes

Há um pico no terceiro flash e a partir daí, há um pico ao fim de cada 4 flashes.
Os picos são cada vez menores ao longo da experiência
Structure of prosthetic groups of b- and c-type cytochromes. The protoheme group
(also called protoporphyrin IX) is found in b-type cytochromes, the heme c group in c-
type cytochromes. The heme c group is covalently attached to the protein by
thioether linkages with two cysteine residues in the protein; the protoheme group is
not covalently attached to the protein. The Fe ion is in the 2+ oxidation state in
reduced cytochromes and in the 3+ oxidation state in oxidized cytochromes.
A model for the organization of electron carriers in PS-I. (From Schubert
et al. 1997.)
The model of Boyer's binding change mechanism. The α and β subunits
configure three nucleotide binding sites: O, which provides the early binding site
for ADP and inorganic phosphate, L, to which the ADP and inorganic phosphate
bind after migrating from O, and T, which tightly binds ATP. Energy ensuing from
the movement of protons from the chloroplast lumen to the stroma drives the
rotation of the γ subunit of CF1, and the interconversion of the binding sites and
the release of an ATP molecule. (From Malkin and Niyogi 2000, after Cross and
Duncan 1996.)
Figure 9.6.B The global distribution of the major sources of fossil fuel
combustion today that leads to increases in atmospheric CO2 levels.
 Ciclo de Calvin-Benson
Melvin Calvin - Nobel da Química (1961)
Nascimento: 8 de abril de 1911, Saint
Paul, Minnesota, EUA
Falecimento: 8 de janeiro de
1997, Berkeley, Califórnia, EUA
Educação: Universidade de
Minnesota (1935), Michigan Technological
University (1931)
Prêmios: Prêmio Nobel de Química, Medalha
Priestley, Medalha Davy,Medalha Nacional de Ciências -
Andrew Benson
Nascimento: 24 de setembro de
1917,Modesto, Califórnia, EUA
Falecimento: 16 de janeiro de 2015, La
Jolla, San Diego, Califórnia, EUA
Educação: Instituto de Tecnologia da
Califórnia, Universidade da Califórnia em
Berkeley
A model for the structure of rubisco in chloroplasts from higher plants.
Rubisco consists of 8 large (L) and 8 small (S) subunits arranged as 4
dimers. Small subunits are shown in red (only four of the small subunits
are seen), large subunits are shown in blue and green, in order to show
the boundaries of the dimers. (From Malkin and Niyogi 2000.)
LUZ
Rubisco ativase
Sistema tioredoxina
Solubility of CO2 and O2 as a function of temperature.
The three variants of the C4
photosynthetic carbon cycle.
The variants differ principally in
(1) the nature of the four-carbon
acid (malate or aspartate)
transported into the bundle
sheath cell and of the three-
carbon acid (pyruvate or
alanine) returned to the
mesophyll cell and (2) the
nature of the enzyme that
catalyzes the decarboxylation
step in the bundle sheath cell.
The three variants are named
after the enzymes that catalyze
the decarboxylation reactions.
Representatives of each variant
include maize, crabgrass,
sugarcane, sorghum (NADP
malic enzyme); pigweed, millet
(NAD malic enzyme); guinea
grass (phosphoenolpyruvate
carboxykinase).
FOTORRESPIRAÇÃO
a) Descarboxilação Via Enzima Málica Dependente de NADP+ (EM-NADP+)
b) Descarboxilação Via Enzima Málica Dependente de NAD+ (EM-NAD+)
c) Descarboxilação via PEP-Carboxicinase
 A map of the geographical abundances of C3 and C4 grasses in the
savannas and grasslands of the world. Courtesy of Ehleringer,
Cerling, and Dearing (2005).
Calculations of the
percentage of the
grass species that
have
C4photosynthesis in
different states of
the western United
States. Note that the
presences of
summer rain
influences C4
abundance. C4 plant
tend to occur in
warm, summer wet
regions. Courtesty of
Ehleringer, Cerling,
and Dearing (2005).
CAM

PEP –Ser OP

PEP- Ser OH
FATORES QUE AFETAM A FOTOSSINTESE
Intensidade luminosa
Temperatura
Efeito de temperatura na fotossíntese de plantas C4 (Tidestromia
oblongifolia) e C3 (Atriplex glabriuscula).
Concentração de CO2
Corn. Courtesy of the Univesity of Tennessee, College of Agricultural
Sciences and Natural Resources:
http://www.utextension.utk.edu/fieldCrops/corn/index.htm
Sugar cane. Courtesy of USGS, South Florida
Virtual Tour:
http://sofia.usgs.gov/virtual_tour/pgcontrolling.ht
ml
Sorghum. Courtesy of AFRICANCROPS.NET:
http://www.africancrops.net/rockefeller/crops/mill
et/pics/kapran-sorghum5a.jpg
 Dicas...
• Importâncias da fotossíntese:

1) Fonte de alimento (200 bilhões de

toneladas de matéria orgânica/ano) que
inicia as cadeias alimentares na
Biosfera.
2) Fonte de O2: sobrevivência dos aeróbicos
3) Existência do O3: proteção contra UV
 Alguns esclarecimentos:
• Plantas realizam fotossíntese somente durante o
DIA (depende da luz) e respiram NOITE e DIA
(independe da luz).
• A intensidade luminosa na qual fotossíntese e
respiração se igualam: Ponto de Compensação
Fótico ou Luminoso.
• Plantas iluminadas abaixo do PCL: mais
consomem do que produzem alimento (morte).
• Plantas iluminadas acima do PCL: mais
produzem do que consomem alimento
(crescem, florescem, acumulam amido).
Modulation of rubisco by rubisco-activase. The combination or the interaction of rubisco (E) with modulators
distributes the enzyme among different states (yellow boxes). The unprotonated form of a specific lysine residue
(E-NH2) reacts with CO2 releasing a proton and forming the carbamate (E-NH-CO2-) that is stabilized by the
cofactor Mg2+ (E-NH-CO2–. Mg2+). At this stage (blue box), rubisco is catalytically competent and thereby binds
ribulose-1,5-bisphosphate (RuBP) yielding the ternary complex (E-NH-CO2– . Mg2+ . RuBP) in which the latter is
converted to the enediol form. This entails the incorporation of CO2 or O2 in the active site to generate the
products of carboxylation or oxygenation, respectively. The high affinity of ribulose-1,5-bisphosphate for free
rubisco (E-NH3+) generates the dead-end complex (E-NH3+ . RuBP) thereby displacing the above equilibria to a
catalytically incompetent state. Rubisco-activase (RA) interacts noncovalently with the binary complex and
facilitates the release of ribulose-1,5-bisphosphate (red box) freeing the enzyme for the next carbamylation cycle.
This process requires the concurrent hydrolysis of ATP.

How the Calvin Cycle Was Elucidated
Autoradiograms showing the
labeling of carbon compounds
in the alga Chlorella after
exposure to 14CO2. The time
intervals shown in the figures
indicate the length of exposure
to the radiolabel. At the
indicated time intervals, the
reaction was terminated, by
plunging of the contents into
boiling alcohol. The labeled
compounds in the cell
homogenates were then
separated by paper
chromatography. The heavy
labeling of 3-phosphoglycerate
(PGA) after the shorter
exposure indicates that it is
the first stable intermediate of
the Calvin (reductive pentose
phosphate) cycle. (After
Bassham 1965.)
GF (sucrose) + GF (sucrose) → GFF (kestose) + G (glucose)
 QUIMIOSSÍNTESE

―Montagem‖ da glicose a partir da energia

química liberada de certas reações
promovidas por algumas bactérias.

Ex: bactérias nitrosas e ferrosas

The value of Φ for a particular process can range from 0 (if that
process is never involved in the decay of the excited state) to 1.0
(if that process always deactivates the excited state). The sum of
the quantum yields of all possible processes is 1.0.
In functional chloroplasts kept in dim light, the quantum yield of
photochemistry is approximately 0.95, the quantum yield of
fluorescence is 0.05 or lower, and the quantum yields of other
processes are negligible. The vast majority of excited chlorophyll
molecules therefore lead to photochemistry.
The quantum yield of formation of the products of
photosynthesis, such as O2, can be measured quite accurately.
In this case the quantum yield is substantially lower than the
value for photochemistry, because several photochemical events
must take place before any O2 molecules form. For O2
production the measured maximum quantum yield is
approximately 0.1, meaning that 10 quanta are absorbed for
each O2 molecule released. The reciprocal of the quantum yield
is called the quantum requirement. The minimum quantum
requirement for O2 evolution is therefore about 10 (see textbook
Figure 7.11). Quantitative measurements of the absorption of
light and the fate of the energy contained in the light are
essential to an understanding of photosynthesis.
Antagonistic effects of light on cytochrome oxidation. Far-red light is very
effective in oxidizing the cytochrome f in the chloroplast. If green light is also
present, some of the cytochrome becomes reduced. The two wavelengths have
opposite effects—hence the term "antagonistic." This experiment is one of the
clearest demonstrations of the existence of two photochemical systems in
photosynthesis: one that reduces cytochrome and one that oxidizes it. This
particular experiment was done with a red alga, in which photosystem II (see
textbook Figure 7.14) is driven best by green light, and photosystem I is driven
best by far-red light. (After Duysens et al. 1961.)
Projected changes in the atmospheric CO2 concentrations under
constrasting emission-control projections. Courtesty of: ACACIA:
http://www.acacia.ucar.edu/images/co2.jpg
Irradiance and fluence rate. Equivalent amounts of collimated light strike a flat
irradiance-type sensor (A) and a spherical sensor (B) that measure fluence
rate. With collimated light, A and B will give the same light readings. When the
light direction is changed 45°, the spherical sensor (D) will measure the same
quantity as in B. In contrast, the flat irradiance sensor (C) will measure an
amount equivalent to the irradiance in A multiplied by cosine of the angle α in
C. (After Björn and Vogelmann 1994.)
A three-axis, triangular presentation of the different gradients influencing
the abundances of different C4 photosynthesis subtypes. Courtesy of
Ehleringer, Cerling, and Dearing (2005).