Types of variables:
Quantitative vs Qualitative
Quantitative: A variable you can measure
Qualitative: A variable you can observe
Independent vs Dependent
Independent: The variable we control
Dependent: The variable we measure/observe; we think it depends on the independent variable
Abbreviated IV and DV
Control group: Group where nothing is being changed
Experimental group: Group where you change the independent variable
Hypotheses:
Not a guess
Proposed explanation for something
Based on what we know
What are some types of intermolecular forces?
Dispersion forces/Van Der Waals forces
Electromagnetic attractions
Dispersion forces
Dipole-dipole forces
Hydrogen bonds
H-bonds in water form between partially positive hydrogen & partially negative oxygen
NOT the term for a covalent bond involving hydrogen
Gives water special properties
Important for structure of macromolecules
Special water properties
High heat capacity
High heat of vaporization
High heat of fusion
Cohesion/adhesion
Most common elements in biological systems: C, H, O, N, P, S (Carbon, Hydrogen, Oxygen, Nitrogen, Phosphorus, Sulfur)
Organic Chemistry: subset of chemistry that deals with organic compounds
Organic: contains carbon & oxygen
Macromolecules
Most compounds fall into 1 of 4 groups of macromolecules
Macromolecule: very large molecule, often with complex properties
Most (but not all) macromolecules = polymers
Polymer: long series of repeated units (monomers)
Making and Breaking Polymers
Forming + breaking down polymers involves removing or adding a water molecule
Make polymers via dehydration synthesis or condensation reaction
Break down polymers via hydrolysis
Classes of Macromolecules
4 main types
Carbohydrates
Basic unit: monosaccharide (aka simple sugar) Ex: glucose
Polymer form: polysaccharide (aka complex sugar/carb)
Contain C, H, O (usually in 1:2:1 ratio)
Carbohydrate Functions
Energy
Glucose main substrate of cellular respiration
Short-term storage
Glycogen in animals
Starch in plants
Structure
Cellulose in plants (cell walls)
Chitin in animals
Signaling/communication
Nucleic acids
Basic unit: nucleotide
Basic structure:
5-carbon sugar
Phosphate group
Nitrogenous group (5 of them)
Thymine (DNA)
Adenine (DNA & RNA)
Cytosine (DNA & RNA)
Guanine (DNA & RNA)
Uracil (RNA)
Polymer: no special term
Contains: C, H, O, N, P
2 main types
DNA
RNA
DNA vs RNA
Both nucleic acids
3 main diffs.
DNA = double stranded, RNA = single-stranded
DNA has thymine, RNA has uracil
DNA's sugar = deoxyribose, RNA'S = ribose
Both present in all living cells
Nucleic Acid Function
Information
Lipids
Contains C, H, O, sometimes P
Largely hydrophobic
Hydrophobic: "water-fearing" doesn't mix well w/ water (other term is hydrophilic "water-loving"
3 main types, each w/ special function
Triglycerides
AKA fats and oils
Main function: long-term energy storage
Glycerol + 3 fatty acids
Saturated vs unsaturated
Saturated fats have no C-C double bonds; tend to be solid, animal fat
Unsaturated fats have C-C double bonds; tend to be liquid, plant fat
Phospholipids
Like triglycerides, but 1 fatty acid replaced w/ phosphate
Changes nature of lipid!
Fatty acids = hydrophobic, phosphates = hydrophilic
Phospholipids want to be both
Will naturally create bilayer (origin of cell membrane)
Function: make up cell membrane
Steroids
Distinctive structure: 4 fused carbon rings
Little functional group additions give properties
Primary function: signaling, regulation (think hormones)
Also plays role in cell membranes
Proteins
Contains C, H, O, N, S
Monomer: amino acids
20 amino acids
4 basic parts
Central carbon w/ hydrogen
Carboxyl group
Amino group
R-group
The 20 amino acids vary in their R-group or sidechain
R-group = functional group
Gives amino acid its properties
Properties determine shape/function of whole protein
R-group interactions
Hydrophobic/hydrophilic interactions
Ionic attractions
H-bonding
Disulfide bridges
All of these give a protein its shape, and the shape of a protein determines its function
Denature: alter interactions/shape of protein... loss/change in function
Polymer: polypeptide
Layers of Structure
Analogy: Layers of structure of a book
1. String of letters (thecatjumpsoverthelazydog)
2. Recognizable subunits: words (cat the jumps over lazy the dog)
3. Repeated structures: sentence (the cat jumps over the lazy dog)
4. Multiple sentences strung together: paragraph
Protein Structure
Primary: sequence of amino acids
What we get from reading genetic code
Doesn't say much abt protein
Secondary: repeated structures (alpha helix & beta sheet)
H-bond interactions between "backbone" of amino acids
2 major structures form:
Alpha helix
Beta sheet
Tells us more abt shape/function than primary structure
Tertiary: 3D shape
Determined by actions between R-groups
Gives us 3D shape/function of the protein
Quaternary: several polypeptides interacting
Some proteins are made of multiple polypeptides
Interaction between polypeptides to form single protein = quaternary structure
Protein Functions:
Structure
Transport
Movement
Signaling/communication (including immune response)
Enzymes: Class of protein that catalyzes reaction
Catalyst: Molecule that lowers activation energy of reaction... not consumed in reaction
Typically named by adding "-ase" to what the enzyme acts on (ex: Lactase breaks down lactose)
Enzyme Function:
Enzymes act on a substrate
Reaction takes place @ active site
Enzymes are specific... tend to catalyze 1 and only 1 reaction
Lock and key model: Idea that only 1 type of molecule is shaped correctly to bind w/ active site
Each has its own building blocks/distinctive properties
Cell Theory
All organisms are made of 1 or more cells
Cell is basic unit of living things
All cells come from existing cells
What is a cell?
Membrane-bound structure
Contains all material necessary for life
All cells
Have DNA
Have a cell membrane
Have sub-cellular structures
Why are cells small?
Pretend cell is a cube (actually not a bad approximation)
Surface area = square
Volume = cube
Cells absorb nutrients across their surface area, but use them in their entire volume
Surface Area:Volume ratio
2 main types of cells
Prokaryotic: archaea and bacteria
No membrane-bound organelles, incl. Nucleus
Always single-celled
Most diverse & numerous group
Eukaryotic: animals, plants, fungi, etc.
Membrane-bound organelles incl. Nucleus
Single or multicellular
Complex cells, moderately diverse
Prokaryotic Cell Structure
Cell membrane (all cells have one)
Nucleoid region where DNA is found
Many ribosomes that produce proteins
Cell wall outside cell membrane; provides protection
Capsule outside cell wall; provides protection & helps cell stick to things
Plasmid: small circular bit of DNA carrying 1 to a few accessory genes
Not part of main chromosome
Flagellum: whip-like extension for movement
Fimbriae: extensions that help cell attach to things
Pilus: for exchanging genetic info
Eukaryotic Cells
Similarities with prokaryotes:
Cell membrane
DNA
Ribosomes
Cytoskeleton
Cell wall
Flagella
Differences
Eukaryotes much bigger
Membrane-bound organelles
Cell Membrane:
Phospholipid bilayer
Also contains
Proteins
Carbohydrates
Steroids (eg. cholesterol)
Fluid-mosaic model
Membrane = fluid, elements move around
Membrane composed of many diff parts
Roles:
Defines/delineates cell
Selectively permeable (controls entry/exit)
Communication w/ other cells
Receiving/responding to signals
Holding cell in place
Nucleus
"Control center" of cell
Contains all cell's DNA
Structure:
Nuclear envelope: double membrane
Nuclear pores: allow/control entry and exit via pore complex
DNA contained in chromosomes, combined w/ protein to make chromatin
Nucleolus
Sub-section of nucleus (not separate by membrane)
Dense region of DNA where ribosomes made
Combines rRNA w/ protein
Cytoplasm
Broad: everything outside the nucleus, including organelles
Often used in place of cytosol, the fluid that makes up most of the cell
Most cellular activity occurs here
Ribosomes
Made of rRNA & protein, no membrane
Make proteins
2 subunits: large & small
2 types
Free: float in cytoplasm, make proteins for use in cell
Bound: attached to Rough ER, make proteins for export
Endomembrane System
Large complex of inter-related organelles that share an evolutionary origin
Carries out
Protein synthesis/modification/transport
Metabolism of lipids
Detoxification
Includes
Nuclear envelope
ER
Golgi
Lysosomes
Transport vesicles
Endoplasmic Reticulum (ER)
Large network of membranes & compartments
2 types w/ distinct function
Rough (RER)
Associated w/ ribosomes
Ribosomes make proteins & release them into ER
RER folds/modifies proteins
Proteins "tagged" for specific destinations
All secreted & most membrane-bound proteins travel thru RER
Smooth (SER)
Series of membrane-bound tubes
Production of lipids
Polysaccharide metabolism
Detoxification
Storage of calcium (in some cells)
Golgi Apparatus
"Warehouse" or "shipping center"
Receives proteins from ER, prepares for export: adds sugar "tags"
Directionality
Cis side towards ER, receives transport vesicles
Trans side towards membrane, sends vesicles away
Lysosomes
Small organelle filled w/ digestive enzymes (hydrolytic)
Roles
Breaks down polymers to harvest monomers
Breaks down old organelles for recycling
Digestion in unicellular organisms
Destruction of pathogens
Apoptosis
Energy Conversion
Cells work w/ 2 main types of energy
Light
Chemical
2 main "currencies"
ATP
Sugars
Cells must be able to convert between energy forms
Mitochondria
Conversion of sugar energy to ATP energy
2 membranes, inner membrane folded into numerous cristae
Site of Krebs cycle & electron transport chain
Contains DNA
Chloroplasts
Found in plant & algal cells
Conversion of light energy to chemical energy
3 membranes, innermost contains chlorophyll
Contains DNA
Endosymbiosis
Chloroplasts & mitochondria originated as individual cells
Ancestral eukaryote engulfed prokaryote, but developed symbiotic relationship
The eukaryote ate the prokaryotic chloroplast/mitochondria, but instead of consuming it, the mitochondria/chloroplast lived inside the eukaryote
Evidence:
Presence of DNA
Presence of prokaryotic ribosomes
Prokaryotic double membrane
Structural Organelles
Cytoskeleton
Centrosomes
Cell wall
Cytoskeleton
Network of protein fibers throughout the cell
Gives cell shape, organizes contents
3 components
Microfilaments (smallest)
Intermediate filaments (medium)
Microtubules (biggest)
Similar structure to flagella/cilia
Centrosomes
Animal cells only
Contains 2 centrioles
Organizes cell division
Cell Wall
Plant, algae, fungi cells
Rigid structure outside cell membrane
Provides structural support & protection
Contains openings: plasmodesmata
Diffusion
Particles = always in motion
What happens if they all move around randomly?
Diffusion: movement from high to low concentration
Osmosis
What if smth can't diffuse?
Reminder: large and/or polar molecules can't cross cell membrane
Water can move thru aquaporins
Osmosis: movement of water across a membrane from low solute concentration to high solute concentration
Hypertonic: more concentrated solution when comparing 2
Hypotonic: more dilute solution when comparing 2
Isotonic: solution w/ same amt. concentration when comparing 2
Osmotic Pressure:
Cells are filled w/:
Salts
Sugar
Proteins
Almost always hypertonic to fresh water
Facilitated Diffusion
The movement of specific molecules across cell membranes thru membrane proteins
When diffusion would take too long/be impossible
Hundreds of diff proteins exist
Active Transport
Moves materials against concentration gradient
Only 1 direction
Requires energy (ATP: Adenosine Triphosphate) bc either pushing smth against concentration gradient or pushing smth big
2 types: Molecular & Endocytosis
Molecular Active Transport
Small molecules & ions
Allows concentration of substances in a particular location
Ion pumps
Endocytosis & Exocytosis
Large molecules & even solid clumps of materials
Endocytosis: Material engulfed by cell thru pockets of cell membrane
Pocket eventually forms vesicle in cytoplasm
Phagocytosis
Large particles
Extensions of cytoplasm surrounds & engulfs particle
Requires considerable amt of energy
Type of endocytosis
Pinocytosis: Tiny pockets form along cell membrane, fill w/ liquid, & pinch off to form vesicles in cell
Exocytosis: vesicle membrane fuses w/ cell membrane, forcing its contents out of the cell
Many cells also release large amts of materials from the cell
Terminology
Metabolism: sum of all reactions occuring in cell
Catabolic: breaking down
Anabolic: building up
Autotroph: organism capable of carbon fixation
Heterotroph: organism incapable of carbon fixation
Carbon fixation: inorganic carbon -> organic carbon
Energy
Comes in many forms
2 main forms cells use
Chemical
Light
2 main "currencies"
Glucose: long term, stable
ATP: shorter term, easy to break
ATP:
3 parts
Adenosine
Ribose
3 phosphates
Energy stored in bonds
ATP & Energy:
ATP uses centers around phosphates
Hydrolysis ... cuts off final phosphate, yielding ADP and energy
ADP: Adenosine Diphosphate
Condensation reaction ... adds phosphate & energy ... "charges" ATP
Bonds & Energy
Breaking bonds takes energy
Hydrolysis of ATP releases energy
Electrons
Phosphates are dense in electrons
Electrons repel each other
Electrons in ATP are in high energy state
After hydrolysis, electron returns to lower energy state, releases energy
Energy cannot be created nor destroyed
Metabolism is all abt energy transfer to do work
Chemical work, eg. digestion
Mechanical work, eg. movement
Transport work, eg. ion pumps
Coupling: endergonic work (requires energy) coupled to exergonic reaction (ATP hydrolysis)
About 99% of energy comes from sun
Overall metabolism: harvest energy from photons, transfer (via intermediates) to do work
Intermediates
Glucose
ATP
NAD+ / NADH ... FAD / FADH2
Mobile electron carriers
Assist in transfer of energy
NADP+ / NADPH in plants
Cellular Respiration
Aerobic cellular respiration has 3 stages
Glycolysis (anaerobic)
Krebs cycle/citric acid cycle (aerobic)
Electron transport chain (aerobic)
Aerobic = requires oxygen; anaerobic = does not require oxygen
Location (in eukaryotes)
Glycolysis in cytosol
Krebs cycle and Electron transport chain in mitochondria
What to keep track of:
For each stage:
Where it takes place and if it needs oxygen
What goes in (type & number)
What comes out (type & number)
Most bookkeeping is "per glucose"
Ask...
What happens to the carbon?
What electron carriers are involved?
Do we get or spend ATP?
Stage 1: Glycolysis
Glyco = sugar, lysis = splitting
1 glucose split into 2 pyruvate
Inputs: Sugar, ATP, NAD+
Outputs: ATP, NADH
Glucose breakdown
Glucose is overall source of energy, but bonds need to be broken
Initial phase requires 2 ATP
Adds phosphates
Destabilizes carbon-carbon bonds
Energy-dense electrons harvested, breaks bonds to yield 2 pyruvates
Additional electrons will be harvested in citric acid cycle
ATP Production
Glucose breakdown lowers free energy... the products have lost energy
Energy has to go somewhere
Energy is used to "charge" 4 ATP
Net gain of 2 ATP
NADH Production
Some free energy transferred to make ATP
Some free energy transferred in form of electrons
2 NAD+ EACH are reduced w/ 2 electrons, yields 2 NADH
Electrons hold energy; will be used in electron transport chain
Inputs
1 Glucose
2 NAD+
2 ATP
2 ADP (usually ignored)
Outputs
2 pyruvate
2 NADH
4 ATP (for a NET gain of 2)
Net gain: 2 pyruvate, 2 NADH, 2 ATP
Getting Ready for Krebs
At end of glycolysis, we have 2 pyruvate
Pyruvate still contains a lot of energy'
Pyruvate Oxidation: precursor step to Krebs cycle
Each pyruvate oxidized w/ acetate, combined w/ CoA (coenzyme A) to make acetyl-CoA
1 NAD+ reduced to NADH (2 total)
1CO2 produced (2 total)
Citric Acid Cycle (Overview)
8-step cycle for harvesting electrons
Runs once per pyruvate (2x per glucose)
By end:
Glucose completely oxidized
High-energy electrons transferred
Some energy (ATP) created
Citric Acid Cycle (Carbon)
Carbon enters cycle as 2 acetyl-CoA
CoA "drops off" 2 carbons... combines w/ oxaloacetate to form citric acid
Over series of steps, carbon oxidized
Electrons transferred to form NADH
Oxidized CO2 released
Oxaloacetate regenerated to restart cycle
Citric Acid Cycle (Electrons)
If carbon is oxidized, smth must be reduced
High-energy electrons transferred to electron carriers
3 NADH per cycle
1 FADH2 per cycle
These electron carriers will return in ETC
Citric Acid Cycle (Energy)
Overall, CAC doesn't produce much free energy
Most energy will be used in ETC
Small amt used to make single ATP per cycle
Products (per glucose)
Pyruvate oxidation:
2 NADH
2 CO2
Krebs cycle:
6 NADH
2 FADH2
4 CO2
2 ATP
Grand total (everything so far)
10 NADH
2 FADH2
6 CO2
4 ATP
Electron Transport Chain (Overview)
Electron Transfer
NADH and FADH2 holding high-energy electrons
ETC starts w/ transfer of e- to ETC protein complexes
Electron carriers oxidized back to NAD+ and FAD
Oxidized electron carriers return to glycolysis/Krebs cycle
Using Electrons
As electrons travel thru ETC they transition from high to low energy
Reminder: energy must go somewhere
Energy used to do work: pump hydrogen ions across membrane
Forms proton gradient
Fate of the Electrons
At end of ETC, electrons have lost most of their energy
What has been pulling them thru the chain?
Oxygen: highly electronegative, attracts electrons
At end of chain, oxygen is reduced, reacts with hydrogen to form water
Without oxygen, nothing to pull electrons through chain
Protons & ATP
Action of ETC proteins leads to proton gradient
High concentration of H+ in intermembrane space
Protons wanna flow back into matrix
1 way back: ATP synthase
ATP Synthase
Large, complex protein embedded in inner membrane
Contains channel allowing proton flow
Proton flow causes enzyme to spin
Energy of spinning used to catalyze ATP formation
Oxidative phosphorylation
ATP Formation
Hydrogens flowing down concentration gradient provide energy for ATP formation
At maximum, respiration leads to formation of 36-38 ATP per glucose
In reality, closer to 30 net ATP per glucose
Leaky membranes
Some energy spent moving things into mitochondrial matrix
Glycolysis, oxygen, & ATP
Observation 1: glycolysis is anaerobic process
Observation 2: if ETC stops, glycolysis will also stop
NADH oxidation
If the ETC stops, NADH will not be oxidized back into NAD+
NAD+ is oxidizing agent in glycolysis
W/out NAD+, nothing to keep glycolysis going
Fermentation: process of allowing NADH to allow additional glycolysis
Caveat: fermentation does NOT generate ATP
Step after glycolysis to regenerate NAD+ in anaerobic conditions
2 types:
Lactic acid fermentation
After glycolysis, pyruvate can be reduced instead of oxidized
1 way: react w/ NADH, picks up e-s (and hydrogen) to become lactate
NAD+ regenerated for glycolysis
Lactate = waste product
source of burning sensation in intense exercise
Alcohol/ethanol fermentation
Another form of regenerating NAD+
2-step process
Produces CO2 and ethanol
Commercially important
Alcohols (ex: beer, wine)
Bread (CO2 makes bread rise)
Fermented vegetables (ex: sauerkraut, kimchi)
Yogurts
Some condiments (ex: soy sauce)
Overall Recap
Glucose has been completely oxidized (stripped of high energy electrons)
Some energy used to make ATP
Most energy used to power ETC, via NADH & FADH2
ETC creates proton gradient that powers ATP synthase
Roughly 30 ATP generated per molecule of glucose
Intro
Photosynthesis: conversion of light energy to chemical energy
Inputs: light & water
Outputs: glucose
Carbon reduced using energy harvested from photons (opposite of cellular respiration)
Equation:
6CO2 + 6H2O + energy -> C6H12O6 + 6O2
Photosynthesis Overview:
2 parts: Light-independent & light-dependent reactions
Light-dependent reactions (LDR):
Harvest light energy to energize electrons
Electrons used to pump hydrogens and reduce NADP+
Creates ATP & NADPH
Light-independent reactions (LIR)
Also called Calvin cycle or dark reactions
Uses energy from ATP & NADPH to reduce carbon
End product: useable carbon
Carbon fixation
Light, Energy, & Pigments
Light form of electromagnetic energy
Basic unit: photon
All photons carry energy
Shorter wavelength = higher energy
Light absorbed by pigments
Chlorophyll: primary photosynthetic pigment
Absorbs primarily red & blue light
Assisted by accessory pigments
Comes in many forms (a, b, c, d, e, & f so far)
Arranged in light-harvesting complexes like solar panels
Works by absorbing photon, using energy to excite electron
Excited electron is at higher energy level
Chlorophyll passes electron to other molecules, becomes oxidized
Electrons will function much like those in ETC of respiration
Chloroplasts
All of photosynthesis takes place in chloroplasts
Structure:
Triple membrane
Innermost membrane: thylakoid stacked into grana
Site of chlorophyll
Lumen: innermost space in thylakoids
Stroma: space between thylakoid & inner membrane
LDR occur across thylakoid membrane
LIR occur in stroma
Photosynthesis part 1
1st part of photosynthesis: light-dependent reactions
Inputs: water & light
Outputs: oxygen (waste), ATP, NADPH
Overall process:
Photon excites electron, energy used to do work
Excited electron replaced w/ electron from water
ETC: proton pumps create proton gradient
ATP synthase harvests proton gradient to make ATP
Electron used to reduce NADP+ to NADPH
Light-Dependent Reactions: Harvesting Light
Process starts w/ chlorophylls absorbing photons in Photosystem II (PSII)
Charged electrons passed to electron transport proteins
Chlorophyll now down an electron, regains it from water
Water is oxidized, creating hydrogen & oxygen
Source of the oxygen gas we breathe
Light-Dependent Reactions: Electron Transport
Excited electrons contain energy, can be used to do work
Pass thru ETC, go from high to low energy
Energy has to go somewhere
Proton pumps create proton gradient
Functionally just like ETC in cell respiration
Light-Dependent Reactions: Harvesting More Light
While ETC is happening, Photosystem I (PSI) also absorbing light
Excited chlorophyll passes electron to 2 possible destinations
Back to ETC
NADP+ to make NADPH
Destination of electron depends on needs for ATP vs NADPH
Lost electron replaced w/ low-energy electron leaving ETC
Light-Dependent Reactions: Summary
Light harvested to energize electrons in PSII
Electrons used to do work: pump protons
Electrons eventually recharged in PSI, can do more work or be transferred to make NADPH
Electrons lost from chlorophyll replaced with electrons from water; creates oxygen
Protons pumped by electrons flow thru ATP synthase to make ATP
Overall products: oxygen, ATP, NADPH
Oxygen = waste
ATP & NADPH will be used in Calvin cycle
Photosynthesis Part II: the Calvin cycle
LDR converts light energy to chemical energy
LIR converts short-term chemical energy to longer-term storage
Occurs in stroma
Summary: uses chemical energy from LDR to fix and reduce carbon
Calvin Cycle: Summary
Aka light-independent reactions, dark reactions, etc.
Do not directly depend on light
Inputs:
CO2
NADPH
ATP
Main product: sugar
3 phases
Carbon fixation
Sugar reduction
Regeneration of RuBP
Calvin Cycle: Carbon Fixation
Cycle starts w/ addition of carbon to 5-carbon sugar called ribulose biphosphate (RuBP)
Performed by enzyme called ribulose-1, 5-biphosphate carboxylase-oxygenase aka RuBisCo
Each reaction represents 1 new fixed atom of carbon (so a glucose requires 6 turns of the cycle)
Calvin Cycle: Sugar Reduction
Remember: glucose is a reduced form of carbon
To hold energy, carbon must gain electrons/be reduced
Phase 2 of Calvin Cycle reacts NADPH and ATP with sugar to transfer high-energy electrons
Eventual product after 3 cycles: 1 glyceraldehyde-3-phosphate (G3P)
Calvin Cycle: Regeneration of RuBP
Reminder: cycles end w/ starting material
Remaining carbons will be rearranged to regenerate original RuBP
Requires spending some ATP
Overall, for every 1 G3P produced, 5 3-carbon sugars are rearranged into 3 5-carbon RuBP
Water & Gas
Plants in arid places can have multiple adaptations
1 adaptation: closing stomata
Pro: no losing water
Con: no exchanging gases
RuBisCo
Carboxylase-oxygenase
If there's too much oxygen sugars get oxidized, not reduced
Helping Rubisco
RuBisCo can react w/ oxygen or CO2
Problem: oxygen and CO2 build up relative to each other
Can be solved by separating LDR. & LIR
2 ways:
C4 plants: separate reactions spatially
CAM plants: separate reactions temporally
C4 Pathway
Way to separate oxygen production & carbon fixation
2 distinct parts of leaves
Mesophyll cells: LDR and initial carbon fixation
Bundle-sheath cells: Calvin Cycle
Carbon is fixed TWICE
1st in mesophyll by PEP carboxylase
Carried by oxaloacetate to bundle
Second time in bundle-sheath by Rubisco
CO2 is concentrated near Rubisco, increases efficiency
CAM pathway
Crassulacean Acid Metabolism
Carbon also fixed twice
Stomata open at night, carbon fixed as malic acid, stored in vacuoles
Respiration lowers oxygen levels
During day, stomata closed, carbon released from malic acid for Calvin cycle
Fixation reactions separated over time
Less efficient, but better at saving water
Controls on Photosynthesis
Water availability, but usually not limiting
CO2 availability
Light
Temperature