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(A* Graded) Summary notes on different forms of energy transfers topic from unit 5. Feature diagrams and concise, informative and detailed notes on: 3.5.1 photosynthesis • The light-dependent reaction • The light-independent reaction 3.5.2 - respiration • Glycolysis • Link reaction and Krebs cycle • Oxidative phosphorylation • Anaerobic respiration 3.5.3 - energy and ecosystems • Food chains and energy transfer • Energy transfer and productivity • Nutrient cycles • fertilisers
Typology: Study notes
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Photoautotrophs utilise photosynthesis to convert energy supplied by absorption of visible light photons from the sun to
chemical energy in the form of intracellular ATP. ATP supplies used to drive metabolic processes through phosphorylating cycles
The net result of photosynthesis is generation of sugars via fixation of atmospheric carbon dioxide
Sugars synthesised are stored for use as substrates used in respiration
Photosynthesis occurs during periods where light incidence on leaf surface is sufficient, plants most photoactive
Cell bodies of mesophyll elongated and contain few other
organelles, maximum volume for occupation by chloroplasts
▪ Large thin + flat leaf
⇢ maximises absorption surface
⇢ minimises diffusion distance for gas exchange
▪ Transparent cuticle/epidermal layer permits passage of sunlight
guard cells control stomatal aperture - responsive to diurnal
change in light intensity
▪ Starch granuales allow storage of carbohydrate - energy reserves
▪ Air spaces in spongey mesophyll - efficient passage in gas phase
▪ Dual thylakoid membrane
Creates compartments between stroma and lumen, provides membrane to cause ion separation
⇢ allows proton gradient to be established across membrane of thylakoid between lumen and stroma
embedded complexes form redox chain
⇢ transfer of electrons from NADH to O 2
, shuttling down series with higher reduction potentials
⇢ Small steps, less exothermic energy loss .energy instead stores in proton gradient
Membrane pumps
▪ High phytopigment content, maximal light capture
contain own biosynthetic machinery
Distinct DNA and ribosomes
⇢ fast manufacture of proteins
⇢ Allows surplus of RuBIsCo to be generated
highly bioavailable enzyme allows fixation via
more energetically favourable route
Photosynthetic tissue specialised to maximise efficiency of chloroplast function. Photosynthetic tissues dominant upper leaf
surfaces, Highest chloroplast abundances in parenchyma cells in the upper mesophyll layer found in aerial parts of plant where
sunlight exposure is maximised
PIGMENTS
Aggregates of photosensitive pigments contained in light harvesting complexes of PSII absorb and redirect energy
towards reaction centres which contain the final chlorophyll a which undergoes excitation.
Broad class of accessory pigments which encompasses range
of colours and absorb in blue-green range of spectrum.
400 - 500 nm range absorbance maxima
⇢ Non overlapping absorption spectra broaden range
absorbable in photosynthesis
Depends on the interplay of two photosystems, linked by common intermediates.
Electrons are given up in the oxidation stage (PSII) and participate in series of reductions in the electron transport chain,
shuttle down membrane before acceptance at the PSI
OUTCOME Convert light energy into chemical energy Use stored chemical energy to fix CO 2
and create a
product that can be converted into glucose
SITE thylakoids Stroma
INPUT Sunlight, H 2
2
PHOTOABSORPTION
Initial absorption of photons occurs in antenna complex
Photosynthetic pigments in granal membrane-embedded protein matrices absorb and funnel energy from incident sunlight
Cascade through antennae complex between pigments terminates at reaction centre chlorophyll-a,
A sufficient quanta of energy is absorbed to cause photoinduced electron excitation
⇢ Electron enters redox series.
Each antenna complex has 250 - 400
accessory pigment molecules.
Diversity of accessory pigments with different
absorption optima broaden the range of
absorbable wavelengths.
In vascular plants accessory pigments include
chlorophylls, xanthophylls, and carotenes.
Chlorophylls constitute primary class of light-absorbing pigment present in green plants
Absorption maxima occur in regions corresponding to and red
Photoionization occurs when chlorophyll a molecules at the reaction centre is raised to higher energy state
⇢ phytolysis of water supplies electrons which reduce chlorophyll and restore it to ground state.
Photoinduced Electron transfer - transfer of excited electron from donor to acceptor generates charge separation.
ETC is comprised by a series of transmembrane embedded protein complexes, involves reduction of carriers at
successively lower energy levels,
electron transfer is exergonic, energy release during transfer exploited to drive
photolysis catalysed by PSII (H 2
O photodissociates into H+ released into the thylakoid lumen + O 2
Shuttling of protons into thylakoid space via active transport to generate gradient
⇢ movement of proton back across membrane coupled with atp synthase pumps
Terminal acceptance of electrons in the PSI by activated carrier NADP+, NADPH formed
2e → NADPH + H
⇢ NADPH exported to stroma for ue in LIR
light + 2H
2
O + 2ADP + 2P
i
O
2
G3P ⁐ RuBP
ANTENNAE COMPLEX - 4 PHOTONS
Light independent stage concludes with the biosynthesis of organic carbohydrates via fixation of atmospheric CO
LIR involves 3 intermediate phases - carboxylation, reduction and RuBP regeneration.
no direct conversion of CO 2
15 out of 18 carbons from the 6 G3P used to regenerate RuBP
1 G3P exported to cytosol for sugar synthesis
⇢ 1 glucose molecule made from 2 triose sugar molecules requires 6 cycles.
Compartmentalization, plasma membrane allows distinct environments to be maintained between cytoplasm and stroma.
⇢ maintains specific enzymatic and chemical composition
3 CO 2
PSI - FINAL ELECTRON ACCEPTANCE
NADP
2H
2e → NADPH + H
G3P ⁐ RuBP
PSII- PHOTOLYSIS
2
2
6
12
6
LIGHT
Dependency on sun arises from photolytic function of light.
The light compensation point defines when carbon dioxide intake balances oxidation of fixed carbon compounds during cellular respiration.
⇢ net carbon dioxide assimilation is zero
TEMPERATURE
Temperature increases within the range of 10-35 °C
have positive effect on rate of photosynthesis due to
particle kinetics.
When temperature increases in range between
minimum to optimum ( 0 - 25 °C), rate doubles
every 10° increase.
▪
Above optimum rate exhibits continued
increase for short duration but later declines as
the temperature cause enzymes to denature,
loss of function
▪
Temperatures too far below optimum correspond to
kinetic energy content of system being too low to
allow enzymes to function at appreciable enough rate
to match sugar consumption
CO 2
consumption in darkness when plant is not photoactive
▪ Pigment density
▪ Type of pigment
▪ Leaf anatomy (stomata, surface area)
accumulation of carbohydrates ▪
▪ hormones
rate increases
linearly as a function
of light intensity.
CO 2
constitutes about 0.041% of atmosphere,
when not under artificially controlled conditions
theoretical maximum rate for photosynthesis
often never achieved due to insufficient CO 2
intense illumination results in radiation
damage to photosynthetic apparatus.
CO 2
influx = respiratory consumption rate
The rate limiting factor determines f the maximum rate at
which a reaction can occur. When a metabolic process is dependent
on multiple factors, rate is limited by the variable nearest its minimum
value or experiences the greatest insufficiency
ENVIRONMENTAL
▪ light
▪ carbon dioxide
▪ Temperature
▪ Water/ion absorption
▪ Atmospheric pollutant
rate stagnates and further increase in light
intensity fails to increase rate
INTERNAL
Light saturation point occurs at light intensity
of around 10% of full sunlight (10,800 lux)
2
Pyruvate reduced to ethanol + CO 2
▪ pathway restricted to prokaryotes
Alcohol dehydrogenases catalyse
alcoholic fermentation
Pyruvate reduced to lactate
Mechanism employed in respiring tissues during strenuous exercise
⇢ causes lactic acid build up in tissue
pH in the cytoplasm quickly drops when hydrogen ions accumulate in
the muscle, eventually inhibiting glycolytic enzymes
⇢ process is unsustainable, suitable only for short term energy
requirements
NAD+ REGENERATED, lactate broken down by liver back into glucose
Net conversion of glucose to pyruvic acid. Yield per glucose:
▪ 2 pyruvate molecules
sugars enter glycolysis pathway via 2 paths:
Glycogenolysis – glycogen stores expended
2 pyruvate molecules formed when G3P
undergoes oxidation through losing electrons
to oxidising agent NAD+
diphosphorylation of glucose to 6c intermediate
fructose-1,6-bisphosphate formed
Phosphate hexose ⇢ glyceraldehyde- 3 - phosphate
Coenzyme regeneration via electron donation to
acceptor molecule
Acceptor molecule differs depending on oxygen
availability. If no oxygen is present then NAD
regeneration necessitates consumption of pyruvate.
g
2e
Pyruvate shuttled to
mitochondria,
converted to acetyl-
coA for use in the
citric acid cycle
The low-yield reaction replenishes cellular NAD+ supplies and phosphorylates 2 ADP
In mammals, greater quantity of ATP only is accessible through pyruvate entering the Krebs cycle in presence of O 2
Cytosolic pathway that constitutes initial stage of aerobic
respiration and principle ATP generation in anaerobic conditions
3.5.2 - GLYCOLYSIS
Pyruvate entering from glycolytic pathway converted to acetyl-coA via dehydrogenase
Intermediary conversion, no net ATP generation during this stage.
As glycolysis splits glucose into two pyruvate molecules, link reaction occurs twice per molecule of glucose..
pyruvate loses COO- group (decarboxylation) via oxidation (-CO 2
reduction of coenzyme NAD
pyruvate ⇢ e-
Citrate (6C) compound undergoes decarboxylation twice to 4C,
yielding 2 CO2 molecules.
▪ dehydrogenases catalysing decarboxylation NAD+ linked
multiple oxidation reactions result in reduction of coenzymes
⇢ deposit electrons into electron transport chain
FAD (flavin adenine dinucleotide)
Dinucleotide coenzyme which functions as an electron
donor or acceptor, cycling between 4 redox states.
TCA - FAD accepts 2 electrons, reduced to FADH
FADH 2 then reverts to FAD, sending its two high-
energy electrons through the electron transport chain.
energy in FADH 2
is enough to produce 1.
equivalents of ATP via oxidative phosphorylation
▪
OXIDATION
NAD (nicotinamide adenine dinucleotide)
2H
2e
NADH + H
glucose → 2 pyruvate
CoA + NAD
→ Acetyl-CoA + CO 2
2
Coenzymes function as secondary substrates, either energy currencies themselves (such as
NADH and ATP) or carriers of chemical groups such as coenzyme A
GLYCOLYSIS
LINK
OXIDATION
REDUCTION
is reduced to NADH through
electron acceptance. Electrons contributed into
ETC where they terminally reduce O 2
Dinucleotide coenzyme, continuously cycles between:
KREBS CYCLE (CITRIC ACID CYCLE)
Reaction series occuring within the mitochondrial matrix providing
reduced forms of coenzymes NADH and FADH 2 for participation in
later stages of respiratory chain (oxidative phosphorylation)
O 2 not used directly as a substrate but still
required to proceed (obligately aerobic)
⇢ functions as terminal electron acceptor
Requires acetyl CoA, generated from
decarboxylation of pyruvate
3.5.2 - RESPIRATION Page 2
Terminal stage in the metabolic pathway, facilitates the controlled release of free energy that was stored in reduced cofactors.
cells release majority of the chemical energy captured by activated carriers at earlier stages.
In eukaryotes process occurs along the inner membrane infoldings of the mitochondria (cristae).
Sequential redox reactions result in free energy release at each stage of an electron transfer cascade. Electrons flow in direction
of compounds with higher reduction potentials
⇢ terminal electron acceptor is molecular oxygen
▪ Greater quantity of energy release per step results in exothermic heat transfer to surroundings.
▪ Energy conserved through release in multiple steps with smaller, more manageable quantities
CHEMIOSMOSIS
ALTERNATE SUBSTRATES
Glycerol is subsequently phosphorylated and converted to triose phosphate which enters the glycolysis pathway.
The fatty acid component is oxidised to acetyl-CoA in the mitochondria and subsequently enters the Krebs cycle.
The free energy liberated at each stage is
used to generate proton motive force.
Reduction of coenzymes deposits protons
which are moved into perimitochondrial
space via active transport, the ion gradient
established creating difference in
transmembrane electrical potential
Diffusion of H+ ions back across the
mitochondrial membrane down gradient
into matrix through transmembrane
protein generates a proton-motive force
exploited by ATP synthases.
ATP synthases couple phosphorylation of ADP return of protons across the membrane down an electrochemical gradient.
The final acceptor of the electrons is elemental oxygen, which becomes reduced to water.
PROTEIN - Excess amino acids are converted into molecules that can enter the pathways of glucose catabolism.
triglycerides ⇢ 3 fatty acids + glycerol
PHOSPHORYLATION
OXIDATION
acetyl CoA
HYDROLYSIS
Energy transfer between trophic levels is inefficient
Approximately 10% of the net production
value at the preceding trophic level retained
food chains rarely extend beyond 5 levels
due to low efficiencies.
Pathways traced by energy through
communities are determined by three energy
transfer efficiencies: consumption, assimilation
and production.
Chemical energy stored in biomass of
organisms at different trophic levels within a
pyramid exhibits a unidirectional decrease as
more transfers occur.
Successive loss as food web complexifies during
transfer between systems.
Loss of energy means chain cannot support many organisms with high nutritive demands, corresponding to fewer
organisms in higher trophic levels and an uneven distribution between number of individuals in species.
Biomass content provides indicator of the quantity of fixed carbon existing within bodies of organic matter. provides
estimate of molecular stock contained within each successive group lower down the energy pyramid.
Sugars not used as substrates are converted into different classes of organic compounds which constutute bulk of organic
matter in carbon-based life forms
Largest contributant to bulk biomass are plants, constitute basal trophic level
Water content of organisms is highly variable, removal of water allows for greater accuracy of comparison.
Sample mass measured and then dehydrated through heating.
Sample heated until 'constant mass', indicating total desaturation.
Once fully dehydrated the final value for dry mass expressed per unit area occupied by plants can be obtained
⇢ Further calorimetric analysis can occur in which the chemical energy in dry biomass can be measured by burning
it and calculating the amount of heat energy released
Measures availability and accessibility of energy for use by next highest trophic level.
Conversion into organic form via photosynthesis driven by incident solar radiation. Total carbon influx corresponds to its
use by respiring autotrophs.
CO 2 efflux by respiration of organisms
Ecosystems form from interactions of biotic and physicochemical components within an environment.
interactions can be quantified in terms of ecological efficiency based on productivity of organisms comprising a community.
100%
10%
1%
0.1%
Energy quantified in temrs of
ecological efficiency (energy making it to the next level) J m
2
dry biomass content (mass of organic matter) (t ha
1
carbon fixed (C m
2
3.5.3 - ENERGY TRANSFER
Initial absorption of photons occurs in antenna complex
Carbon entry into biosphere occurs via fixation of carbon dioxide by primary producers.
Gross primary productivity (GPP) - total amount of energy directly acquired by autotrophic energy harvesting
⇢ Photosynthetic efficiency averages 5.5% in terrrestrial ecosystems.
Of the energy fixed, 20-50% is used in respiration or other biochemical operations within the plant, accounted for through
net primary productivity (NPP) which deducts respiratory and thermodynamic loss
Quantifies generation of biomass through heterotrophic nutrition.
Expresses quantity of new tissue created through assimilation of
energy consumed through food.
Consumption includes:
primary consumption of plant matter (herbivorous/omnivorous)
⇢ herbivorous assimilation efficiencies 20-50%
Higher consumption levels of animals (omnivorous/carnivorous)
⇢ carnivore assimilation efficiency level of 80%
Ecosystem processes involve complex interactions between detrtitus and predatory food webs, nutrient cycling between
trophic cascades maintains flow and allows continuous generation
Autotrophs absorb organic minerals and fix them in usable organic forms for abbsorption by autotrophs through the soil
Nutrients that are not assimilate return to the ecosystem via two main pathways:
▪ Consumers release mineral nutrients via excretion, direction deposition and return
Defecant and unassimilated organic matter from non-predatory death form detritus. mineralisation by microbes
reintroduces nutrients to soil
Minimising respiratory losses in animals reared for farming increases efficiency of biomass conversion through:
▪ Restricting movement, higher stock density in enclosed compact spaces limits respiratory losses
Keeping ambient environment warm to minimise use of energy for body heat
▪ Controlled feeding so animals recieve optimum nutrients for growth
▪ Exclusion of predators limits losses through predatory deaths
Simplification of food webs - removal of competitive species such as weeds which compete for resources. achieved through:
H
PHOTOSYNTHETIC EFFICIENCY LIMITANTS:
▪ Non-absorbable light wavelengths refracted
▪ Reflection by particulate matter e.g dust, clouds
▪ Light incidence on non chlorophyllic region
▪ Climate/ diurnal changes
▪ carbon dioxide concentrations
▪ Resource scarctiy
The factors that limit terrestrial primary productivity are
solar energy (and particularly its inefficient use by plants),
water and temperature.
Differences amongst biomes, depends on proportion of
the biomass photosynthetically active
fragmentation of plant and animal debris, saprophagous
microorganisms complete decomposition and mineralization
▪ responsible for clearance of organic waste
⇢ In the absence of detritivores, detritus would accumulate,
creating a bottleneck in nutrient cycling and inhibiting further
primary production
DECOMPOSERS (saprobionts)
Saphrophytic organism participate in
extracellular digestion
▪ enzyme secretion through cell membrane
occurs external to a digestive tract, absorption
and metabolism at molecular level
terrestrial : earthworms
aquatic : sea cucumbers, crabs
Energy is lost during conversion into new animal
mass, only fraction of the material ingested (I)
absorbed and assimilated contributing to growth of
organism + reproductive activity.
Most used for respiration (Rh), remainder defecated
Energy loss in secondary production due to:
▪ Inedible parts of organism not consumed
▪ Indigestible parts excreted as undigested waste
⇢ faecal matter replenishes nutrient supply
through saphrophytic nutrition
Decomposition of dead organic matter by microorganisms returns constituent
molecules in the form of detritus to basal pool of an ecosystem
NPP = GPP - R a
Interconversion between allotropes of nitrogen as it circulates between the atmosphere, terrestrial and marine ecosystems.
Plants must secure nitrogen in its fixed form, which requires it to be incorporation in compounds
atmosphere is composed of 79% nitrogen gas N 2
, however this major nitrogen reservoir is not available for use in
biological systems, leading to resource scarcity in many ecosystems.
⇢ N2 (g) bond enthalpy is high, meaning low reactivity and high energy requirements associated with fixation
ALLOTROPE FORMULA PRESENCE IN NATURE
Diatomic nitrogen N 2
Air
Ammonia NH 3
urea, rainwater, manure,
rotting plants
Ammonium NH 4
fertile soil + seawater
Nitrite NO 2
Nitrate NO 3
sodium nitrate or in
mineral forms
Nitrous oxide N 2
O gas emissions
Nitric oxide NO thunderstorms
Nitrogen fixing bacteria and
root nodule symbionts in
association with plants fix
nitrogen and cooperate with
host to supply nitrogen
Ecosystems recycling occurs locally through interconversion of mineral nutrients during biomass transfer in addition to
larger scale participation in a global system of inputs and outputs where matter is exchanged in biogeochemical cycles.
PROCESS conversion of inert nitrogen gas present in the
atmosphere into nitrogenous compounds capable of entering into
terrestrial and aquatic ecosystems from their deposit forms.
Conversion of nitrogen between its various allotropes
#predominantly results from microbial activity, either
in chemoautotrophic energy acquisition or to access
nitrogen required for their growth.
2
formation is catalysed by lightning which provides
sufficient energy to overcome high bond enthalpy.
3 ,
enters soil via precipitation
conversion of H 2
2
into
3
under high pressures
and temperatures in Haber
process. Used in fertilizer
formation
Diazotrophic microorganisms associate with plant rhizosphere to form nodules which
increase surface area of roots. Nitrogenase enzymes fix N 2
into absorbable form.
Association is mutual, symbiosis provides the diazotroph with a steady influx of carbon
sources + regulated oxygen supply tightly bound by leghaemoglobin
RHIZOBIA associate with legume species, infecting root nodules.
MYCORRHIZAE are fungi colonisers of plant roots, form arbuscular tubules, hyphal
extensions increase surface area for uptake of water + minerals
3.5.4 - NUTRIENT CYCLE
AMMONIFICATION
PROCESS conversion of nitrogenous waste from decomposing
organic matter (dead animals/ faeces) into ammonium
Saprobionts mineralize waste via aerobic or anaerobic
decomposition, proteolytic enzymes catalyze formation of
inorganic NH 4
⇢ ammonium salt compounds form in soil + aqueous reserves,
forming deposits which can be nitrified
4
PROCESS conversion of ammonium/ammonia to nitrites
via oxidation, undergo further oxidation to form nitrates.
▪ process utilized by chemoautotrophs
Oxygen required in ammonium and nitrite oxidation,
both nitrifying and nitrite-oxidizing bacteria are aerobes.
3
2
(⇢ absorption)
NITRIFICATION
DENITRIFICATION
PROCESS anaerobic denitrifying bacteria facilitate
conversion of ions in soil into gaseous nitrogen (N 2
▪ returns bioavailable nitrogen to atmosphere,
▪ typically occurs in anoxic environments
2
unavailable for use as terminal electron acceptor in
electron transport, N 2
substituted in anaerobic respiration
▪ Increased usage in chemical fertilizers in agriculture alters nitrogen cycle
Agricultural and industrial nitrogen inputs currently exceeds inputs from
natural fixation, accounting for an increase in nitrogenous compound
deposition to marine + terrestrial ecosystems (+ 200% since 1600)
SYNTHETIC - ammonia derived compounds
generated through Haber process
NATURAL - compost, animal manure,
harvested minerals, crop rotations
Denitrification can be considered an ecologically beneficial and
detrimental process, depending upon when and where it occurs.
(+)
useful in water-treatment processes, removal of
nitrogen from water supplies prior to discharge into
natural bodies prevents eutrophication
(-)
by-products include NO + N 2 O, greenhouse gases which
contribute to global warming
(-)
decrease soil fertility, removing nitrogen from soil means
loss from basal mineral pool
DEPOSITION - FERTILISER USE
EUTROPHICATION
ECOSYSTEM EFFECTS
▪ Soil acidification
▪ Toxic compounds can accumulate in soils or in animals and pass throughout food chain
▪ Ozone degradation due to NO production, acceleration of global warming
▪ Reduction of species diversity as nitrogen rich soils favour fast-growth plants, leading to out competition of other species
▪ Decreased atmospheric visibility
Involves cycling of phosphoric allotropes through the lithosphere
hydrosphere and biosphere. Phosphorus is predominantly found in
sedimentary deposits or in aqueous dissolved phosphate reserves.
lacks gaseous phase, entry of orthophosphates into soils occurs through:
gradual accumulation over geological time as a consequence of
weathering or uplifting / erosion of rocks.
breakdown of manure, chemical fertilisers, biosolids or
plant/animal debris.
Sedimentation of dissolved particulate phosphorus creates layer
of seabed phosphorus rocks which gradually erode
Necessary for plant growth, deficiency
causes decolouration and stunted growth
Uses of orthophosphates in plants:
▪ Nucleic acids
▪ Buffering agent
▪ Phospholipid bilayer
Tectonic uplift exposes phosphorus-
bearing rocks to weathering, physical
erosion of bare rock contributes
dissolved and particulate phosphorus
to soils, lakes and rivers.
Soils have a finite capacity to bind P. When saturated with P, desorption is accelerated, increasinsg soil
leaching and runoff. As too much phosphate is present for all to be absorbed
▪ Loss through removal of crop, phosphates not returned to soil through decomposition