Ch 1 Biochem: Foundations of Biochemistry - Comprehensive Notes

1. Cellular Foundations

  • Life on Earth arose around 4 billion years ago with simple microorganisms that:

    • extracted energy from chemical compounds and sunlight

    • used that energy to synthesize biomolecules from Earth's surface elements and compounds

  • Principle 1: Cells are the fundamental unit of life

    • cells vary in complexity and specialization but share remarkable similarities

    • examples include single-celled bacteria to highly specialized cells in multicellular organisms

  • Principle 2: Cells use a relatively small set of carbon-based metabolites

    • these metabolites build polymeric machines, supramolecular structures, and information repositories

    • the chemical structure of these components defines cellular function

    • the collection of molecules carries out a program, whose end result is replication of the program and self-perpetuation of the molecule collection (i.e., life)

  • Principle 3: Living organisms exist in a dynamic steady state, never at equilibrium

    • follow the laws of thermodynamics by extracting energy from surroundings to maintain homeostasis and do useful work

    • essentially all cellular energy comes from electron flow, driven by sunlight or metabolic redox reactions

  • Principle 4: Cells can precisely self-replicate and self-assemble using genome-stored chemical information

    • example: a single bacterial cell in sterile nutrient medium can yield ~10^9 identical daughter cells in 24 hours; each cell faithfully copies the original via genetic information

    • on a larger scale, vertebrate progeny resemble parents due to inheritance of parental genes

  • Principle 5: Living organisms change over time by gradual evolution

    • results in enormous diversity related through shared ancestry

    • molecular evidence is seen in the similarity of gene sequences and protein structures

  • What is biochemistry?

    • biochemistry = describes the structures, mechanisms, and chemical processes shared by all organisms

  • 1.1 Cellular Foundations

    • Cells are the structural and functional units of all living organisms

  • The Plasma Membrane

    • defines the periphery of the cell

    • composed of lipid and protein molecules

    • thin, flexible, hydrophobic barrier around the cell

    • contains embedded transport proteins, receptor proteins, and membrane enzymes

  • Cytoplasm Contains Cytosol and Suspended Particles

    • cytoplasm = internal volume enclosed by the plasma membrane

    • composed of cytosol (an aqueous solution) plus suspended particles (mitochondria, chloroplasts, ribosomes, proteasomes, etc.)

    • cytosol = highly concentrated solution containing enzymes, RNA, amino acids, nucleotides, metabolites, coenzymes, and inorganic ions

  • Clicker Question 1: Cytosol does not contain which? A) enzymes and RNA molecules that encode them; B) membranous organelles (e.g., mitochondria); C) inorganic ions (K+, Na+, Mg2+); D) intermediates in biosynthetic/degradative pathways

  • Answer: B (membranous organelles sediment when cytoplasm is centrifuged at 150,000 g; the supernatant is the cytosol)

  • The Nucleoid or Nucleus of a Cell Stores the Genome

    • genome = complete set of genes, composed of DNA

    • bacteria/archaea (prokaryotes) store genome in a nucleoid

    • eukaryotes store genome in a membrane-enclosed nucleus

  • Cellular Dimensions Are Limited by Diffusion

    • typical sizes: animal/plant cells 5–100 μm; unicellular microbes 1–2 μm

    • upper size limit set by transport rate and O2 delivery to all parts of the cell

    • example: human lymphocytes

  • Clicker Question 2: All cells: A) have a plasma membrane. B) have mitochondria. C) are approximately the same size. D) are at least partially aerobic.

  • Answer: A (All cells have a plasma membrane; mitochondria are specific to many but not all cells; size varies; aerobic capacity varies)

  • Clicker Question 2, Response: All cells have a plasma membrane; the membrane defines cell periphery and serves barrier/interaction roles

  • Organisms Belong to Three Distinct Domains of Life

    • Bacteria: inhabit soils, surface waters, and tissues of other organisms

    • Archaea: inhabit extreme environments

    • Eukarya: all eukaryotic organisms; more closely related to archaea than to bacteria

  • Phylogeny of the Three Domains of Life

  • Archaea and Bacteria Subgroups Are Distinguished by Habitats

    • aerobic: plentiful O2; energy from electron transfer to O2

    • anaerobic: no O2; electrons transferred to nitrate, sulfate, or CO2

    • obligate anaerobes die in O2

    • facultative anaerobes can live with or without O2

  • Clicker Question 3: New organism in a salt lake that can live with or without O2 would be best classified as: A) obligate anaerobe in Bacteria; B) obligate anaerobe in Archaea; C) facultative anaerobe in Archaea; D) facultative anaerobe in Bacteria

  • Answer: C (facultative anaerobe in Archaea)

  • Clicker Question 3, Response: Archaea inhabit extreme environments; facultative anaerobes can live with or without O2; thus, facultative anaerobe in Archaea

  • Principle 3 (2 of 5): Reiteration of dynamic steady state; energy from surroundings; electrons flow from light or redox reactions

  • Organisms Differ Widely in Their Sources of Energy and Biosynthetic Precursors

    • phototrophs: trap and use sunlight

    • chemotrophs: derive energy from oxidation of chemical fuels

    • autotrophs: synthesize all biomolecules directly from CO2

    • heterotrophs: require preformed organic nutrients from other organisms

  • Classifying Organisms by Energy Source

  • Clicker Question 4: Humans are which type? A) photoautotrophs; B) chemoautotrophs; C) photoheterotrophs; D) chemoheterotrophs

  • Answer: D (humans are chemoheterotrophs): derive energy from chemical fuels and require organic nutrients from others

  • Clicker Question 4, Response: Humans are chemoheterotrophs; explanation

  • Bacteria and Archaeal Cells Share Features but Differ in Important Ways

    • cell envelope: comprised of plasma membrane, outer membrane, and peptidoglycan (high-molecular-weight polymer)

  • Bacterial and Archaeal Cell Envelopes

    • Gram-positive bacteria: thick peptidoglycan layer outside plasma membrane; no outer membrane; Gram stain positive

    • Gram-negative bacteria: outer membrane is a lipid bilayer

    • Archaea: layer of peptidoglycan or protein provides rigidity

  • Principle 2 (2 of 6): Small metabolite set to build cellular machines; program and self-replication

  • The E. coli Cytoplasm

    • contains ribosomes, enzymes, metabolites, cofactors, inorganic ions

    • nucleoid contains a single, circular DNA molecule

    • plasmids: smaller circular DNA segments that confer toxin/antibiotic resistance

  • Eukaryotic Cells Have Various Membranous Organelles (isolated for study)

    • mitochondria: site of most energy-extracting reactions

    • endoplasmic reticulum and Golgi: synthesize and process lipids and membrane proteins

    • peroxisomes: oxidize very-long-chain fatty acids and detoxify reactive oxygen species

    • lysosomes: digestive enzymes

    • granules/droplets: store nutrients (e.g., starch, fat)

  • Clicker Question 5: Which organelle is not membrane-bound? A) lysosome B) peroxisome C) ribosome D) Golgi complex

  • Answer: C (ribosome is not membrane-bound)

  • Clicker Question 5, Response: Ribosome is not membrane-bound; others are enclosed by membranes

  • Plant Cell Organelles

    • vacuoles: store large quantities of organic acids

    • chloroplasts: site of light-driven ATP synthesis (photosynthesis)

  • Eukaryotic Cell Structure

  • Clicker Question 6: Which organelle is found in both plant and animal cells? A) starch granule B) chloroplast C) mitochondria D) glyoxysome

  • Answer: C (mitochondria)

  • Clicker Question 6, Response: Mitochondria are present in both plant and animal cells

  • Activity: Metabolic Map

    • guided exploration of metabolic pathways; focus on shared pathways among prokaryotes and eukaryotes

  • Activity Instructions (1 of 4)

    • Think of three pathways and determine if they are used by eukaryotes, prokaryotes, or both; discuss in pairs and class

  • Activity Solution (1 of 4)

    • Prokaryotes: Glycolysis, Gluconeogenesis, Citric Acid Cycle, Glycogen metabolism, Urea cycle, Oxidative phosphorylation, etc.; Both: many core pathways

  • Subcellular Fractionation of Tissue

    • first step: gently disrupt cells/tissues to rupture plasma membrane

    • second step: centrifuge the homogenate

    • organelles differ in size and sedimentation rates

  • The Cytoplasm Is Organized by the Cytoskeleton and Is Highly Dynamic

    • cytoskeleton = three-dimensional network of protein filaments in eukaryotic cells

    • components: actin filaments, microtubules, intermediate filaments

    • filaments continually disassemble and reassemble

  • Cytoskeletal Filaments: actin filaments, microtubules, chromosomes, kinetochores, centrosomes, intermediate filaments

  • The Structural Organization of the Cytoplasm

    • endomembrane system segregates specific metabolic processes and provides surfaces for enzyme-catalyzed reactions

    • Exocytosis and endocytosis = transport across membranes; involve membrane fusion/fission; connect cytoplasm with surroundings

  • Clicker Question 7: The cytoskeleton is composed of: A) membranous organelles. B) kinetochores and centrosomes. C) actin filaments, microtubules, and intermediate filaments. D) chromosomes and plasmids.

  • Answer: C; the cytoskeleton comprises actin filaments, microtubules, and intermediate filaments

  • Clicker Question 7, Response: Correct—three general cytoplasmic filament types

  • Principle 2 (3 of 6): Small metabolite set to build cellular machines; life program

  • Cells Build Supramolecular Structures

    • held together by noncovalent interactions: hydrogen bonds, ionic interactions, van der Waals interactions, hydrophobic effect

  • In Vitro Studies May Overlook Important Interactions Among Molecules

    • in vitro = in glass; in vivo = in living organism

    • molecules may behave differently in vivo vs. in vitro

  • Clicker Question 8: Supramolecular complexes (e.g., chromatin) are held together by: A) covalent bonds between monomeric units. B) noncovalent interactions (e.g., hydrogen bonds). C) covalent bonds between macromolecules. D) interactions between cytoskeleton and organelles

  • Answer: B; noncovalent interactions hold supramolecular complexes together

  • Clicker Question 8, Response: Noncovalent interactions drive chromatin organization; monomer covalent bonds join subunits, not the supramolecular assembly

  • 1.2 Chemical Foundations

  • Elements Essential to Animal Life and Health

  • Biomolecules Are Carbon-based Compounds with Functional Groups

    • carbon forms covalent single, double, and triple bonds

  • Geometry of Carbon Bonding

    • carbon atoms adopt a characteristic tetrahedral arrangement for four single bonds

    • free rotation around single bonds; limited rotation about double bonds

  • Common Functional Groups of Biomolecules

  • Additional Functional Groups of Biomolecules

  • Clicker Question 9: Important functional groups in biomolecules include: A) lipids. B) thioesters. C) nucleic acids. D) carbons.

  • Answer: B; thioesters are a notable functional group in biomolecules

  • Clicker Question 9, Response: Thioesters confer specific chemical properties; functional groups define families of compounds

  • Many Biomolecules Are Polyfunctional

  • Cells Contain a Universal Set of Small Molecules

    • central metabolites: common amino acids, nucleotides, sugars and their phosphorylated derivatives, mono-/di-/tricarboxylic acids

    • secondary metabolites: organism-specific

    • metabolome = entire collection of small molecules in a cell under certain conditions

    • metabolomics = systematic characterization of the metabolome under precise conditions

  • Macromolecules Are Major Constituents of Cells

    • macromolecules = polymers with MW > ~5,000 built from simple precursors

    • major classes: proteins, nucleic acids, polysaccharides

    • oligomers = shorter polymers

    • informational macromolecules = proteins, nucleic acids, some oligosaccharides with information-rich subunits

  • Protein Macromolecules

    • proteins are long polymers of amino acids; functions include enzymes, structural components, signal receptors, transporters

    • proteome = all proteins functioning in a cell; proteomics = study of this protein set

  • Clicker Question 10: Proteins are macromolecules because: A) they can noncovalently form large structures. B) they are polymers with MW above ~5,000. C) they can function as enzymes, structural elements, receptors, or transporters. D) they are composed of multiple oligomers.

  • Answer: B; proteins are macromolecules because they are polymers with MW > ~5,000

  • Clicker Question 10, Response: Correct—the definition of macromolecules includes high molecular weight polymers

  • Nucleic Acid Macromolecules

    • DNA and RNA = polymers of nucleotides; store and transmit genetic information; some RNA have structural/catalytic roles

    • genome = entire sequence of a cell's DNA or RNA

    • genomics = characterization of genome structure, function, evolution, and mapping

  • Polysaccharide Macromolecules

    • polysaccharides = polymers of simple sugars; roles in energy storage, cell-wall structure, and intercellular recognition

    • glycome = entire repertoire of carbohydrate-containing molecules

  • Lipid Molecules

    • lipids = water-insoluble hydrocarbon derivatives; roles include membrane structure, energy stores, pigments, intracellular signaling

    • lipidome = lipid-containing molecules in a cell

  • Activity: Classifying Metabolites

    • explore structure of palmitic acid in Mol3D viewer

  • Activity Instructions (2 of 4)

    • Is palmitic acid a carbon-based metabolite? What type of biomolecule is palmitic acid?

  • Activity Solution (2 of 4)

    • Palmitic acid is carbon-based (backbone with 16 carbons) and is a fatty acid, a type of lipid

  • Building Blocks of Biochemistry

  • Major Classes of Biomolecules in E. coli Cells (Table 1-1)

    • Water ~70% of weight; Proteins ~15% (≈3,000 species); DNA ~1% (1–4 molecules); RNA ~6% (>3,000 species);
      Polysaccharides ~3%; Lipids ~2%; Monomeric subunits and intermediates ~2%; Inorganic ions ~1%

    • Note: if all fatty acid permutations are counted, lipid diversity is vast

  • Clicker Question 11: The systematic characterization of the entire collection of small molecules in a given cell under a specific set of conditions is called: A) genomics. B) proteomics. C) lipidomics. D) metabolomics.

  • Answer: D; metabolomics

  • Clicker Question 11, Response: Metabolomics = study of the metabolome under defined conditions

  • Three-Dimensional Structure Is Described by Configuration and Conformation

    • configuration = fixed spatial arrangement of atoms

    • stereoisomers = molecules with same bonds and formula but different arrangement

    • stereospecific = interactions require specific conformations

  • Illustrating Stereochemistry

  • Configurations of Geometric Isomers

    • cis/trans (geometric) isomers differ in substituent arrangement around double bonds

  • Chiral and Achiral Molecules

    • chiral centers = stereocenters with four different substituents

    • a molecule can have 2^n stereoisomers where n = number of chiral centers

  • Clicker Question 12: Molecules that differ in configuration cannot be: A) stereoisomers. B) cis-trans isomers. C) chiral centers that are mirror images of each other. D) chiral centers that can be interchanged by rotation of a single bond.

  • Answer: D; chiral centers that can be interconverted by rotation of a single bond are not distinct stereoisomers

  • Clicker Question 12, Response: Configuration depends on double bonds or chiral centers; rotation around single bonds can interconvert conformers but not alter configuration

  • Enantiomers and Diastereomers

    • enantiomers = mirror-image stereoisomers

    • diastereomers = non-mirror-image stereoisomers

  • Optical Activity of Enantiomers

    • enantiomers have similar reactivity but different optical activity

    • racemic mixture shows no net optical rotation

  • Naming Stereoisomers Using the RS System

    • assign priorities to groups attached to chiral carbon

    • priority order example: —OCH3 > —OH > —NH2 > —COOH > —CHO > —CH2OH > —CH3 > —H

  • Clicker Question 13: Enantiomers: A) are only associated with amino acids. B) can be specific types of diastereomers. C) are always designated either D or L. D) can exist for molecules with more than one chiral carbon.

  • Answer: D; enantiomers can exist for molecules with more than one chiral carbon

  • Clicker Question 13, Response: Enantiomers can arise when there are multiple chiral centers; each can yield distinct stereoisomers

  • Molecular Conformation

    • conformation = spatial arrangement of substituent groups that are free to rotate

  • Principle 2 (4 of 6): Small metabolite set to build polymeric and supramolecular structures

  • Interactions Between Biomolecules Are Stereospecific

  • Biological Systems Can Distinguish Stereoisomers

    • stereospecificity = ability to distinguish stereoisomers

  • Clicker Question 14: The antidepressant Celexa is a racemic mixture of two stereoisomers, but only (S)-citalopram has the therapeutic effect. A stereochemically pure preparation of (S)-citalopram is sold as Lexapro. Which is true? A) Lexapro shows no optical activity. B) Biological systems cannot distinguish the two isomers. C) The effective dose of Lexapro is half that of Celexa. D) Both stereoisomers are biologically active.

  • Answer: C (Lexapro’s active enantiomer allows a lower dose to achieve the therapeutic effect)

  • Clicker Question 14, Response: Only the active enantiomer contributes to efficacy; racemate requires higher total dose for effect

  • 1.3 Physical Foundations

  • Living organisms exist in a dynamic steady state, never at equilibrium with surroundings

    • small molecules, macromolecules, and supramolecular complexes are continually synthesized and broken down

    • steady state requires constant energy input

  • Organisms Transform Energy and Matter from Surroundings

    • system vs. universe definitions; open/closed/isolated systems

    • open systems exchange both energy and matter with surroundings

  • Clicker Question 15: A living organism is a(n): A) isolated system. B) closed system. C) open system. D) universe.

  • Answer: C; open system

  • Clicker Question 15, Response: Open system exchanges energy and matter with surroundings

  • Energy Transformation in Living Organisms

    • first law of thermodynamics: energy is conserved; can change form

  • Extracting Energy from Surroundings

    • photoautotrophs vs chemotrophs

  • Oxidation-Reduction Reactions

    • redox reactions involve transfer of electrons; linked to energy flow in metabolism

    • autotrophs/heterotrophs participate in global O2/CO2 cycles powered by sunlight

  • Creating and Maintaining Order Requires Work and Energy

    • second law: total randomness tends to increase; living systems maintain order by inputting energy

    • entropy S measures randomness/disorder

  • Free Energy, G

    • enthalpy H ≈ heat content

    • free energy G = H − T S

    • in a closed system, G predicts work potential and spontaneity

    • G=HTSG = H - T S

  • Clicker Question 16: Which reflects the total energy change in a chemical reaction, i.e., bonds formed/broken? A) ∆G B) ∆H C) ∆S D) ∆T

  • Answer: B; ∆H

  • Clicker Question 16, Response: Enthalpy change ∆H reflects bond energies and noncovalent interactions broken/formed

  • Free-Energy Change, ∆G

    • riangleG=riangleHTriangleSriangle G = riangle H - T riangle S

    • spontaneous reactions have riangle G < 0

    • for reactions in a closed system, reactions proceed toward equilibrium until ∆G = 0

  • Coupling Reactions

    • endergonic reactions can be driven by coupling to exergonic reactions (e.g., ATP hydrolysis)

    • breaking phosphoanhydride bonds in ATP is highly exergonic

  • Energy Coupling Links Reactions in Biology

    • riangleG=riangleGextrxnextactual=ext(energyavailabletodowork)riangle G = riangle G_{ ext{rxn}}^{ ext{actual}} = ext{(energy available to do work)}

    • in closed systems, reactions proceed spontaneously until equilibrium; in living systems, coupling maintains non-equilibrium states

  • Clicker Question 17: If A → B has ∆G = −14 kJ/mol and C → B has ∆G = +16 kJ/mol, then: A) A→C reversible. B) A→C exergonic. C) A and C can never be at equilibrium. D) A→C is entropically driven.

  • Answer: B; A→C is exergonic (∆G = −30 kJ/mol if combined)

  • Clicker Question 17, Response: Demonstrates that combining reactions can yield a larger exergonic drive

  • Keq and ∆G° as Measures of Spontaneity

    • Keq = ([products]eq)/([reactants]eq) at equilibrium

    • Mass-Action Ratio, Q = [products]/[reactants] at any time; used to gauge distance from equilibrium

    • relation: riangle G = riangle G^ of the reaction? + RT \,

    • standard free-energy change, ∆G°, is the free energy change under standard conditions; actual ∆G depends on concentrations

  • Mass-Action Ratio, Q

    • Q = ([A]^{a} [B]^{b} …)/([C]^{c} [D]^{d} …)

  • Clicker Question 18: ATP breakdown yields ADP and Pi; Keq = 2×10^5 M. With cellular [ATP] = 15 mM, [ADP] = 1.5 mM, [Pi] = 15 mM, is this reaction at equilibrium in living cells?

  • Answer: D; No, because Q ≪ Keq. Calculation: Q = [ADP][Pi]/[ATP] = (1.5 mM)(15 mM)/15 mM = 1.5 mM, far from Keq

  • Clicker Question 18, Response: Q is much smaller than Keq; reaction far from equilibrium in cells

  • Standard Free-Energy Change, ∆G°

    • ∆G actual for a reaction depends on ∆G° and conditions: riangle G = riangle G^\u00b0 + RT \, ext{ln} Q

    • details: A reaction with ∆G° and actual conditions yields current ∆G

  • Clicker Question 19: A reaction will always be thermodynamically spontaneous if: A) ∆H < 0 and ∆S > 0. B) ∆G = 0. C) it is coupled to another reaction. D) ∆G° > 0.

  • Answer: A (and due to coupling as needed); spontaneous when ∆G < 0

  • Clicker Question 19, Response: Spontaneity requires ∆G < 0; A combines negative enthalpy and positive entropy contributions

  • Reactions Do No Work at Equilibrium

  • Reaction Coordinate Diagrams and Enzymes

    • exergonic reactions can be coupled to endergonic reactions to drive processes toward biological work

    • enzymes greatly increase reaction rates by stabilizing transition states

    • Activation energy: riangle G^\u00b0

    • Enzymes do not change the equilibrium, only the rate

  • Clicker Question 20: What function do enzymes perform? A) Increase equilibrium. B) Increase rate but not equilibrium. C) Increase activation energy. D) Make products higher energy than reactants.

  • Answer: B; enzymes increase rate without altering equilibrium

  • Clicker Question 20, Response: Enzymes are biocatalysts; lower activation energy, raise rate; do not affect equilibrium

  • Catabolism and Anabolism

    • Pathways are sequences of consecutive reactions where product of one becomes substrate of next

    • Catabolism: degradative, energy-yielding; drives ATP synthesis and NAD(P)H production

    • Anabolism: synthetic pathways requiring energy input

  • Metabolism

    • metabolism = network of enzyme-catalyzed pathways (both catabolic and anabolic)

    • unity of life: core pathways acting on proteins, fats, sugars, nucleic acids are largely identical across organisms

  • Metabolism Is Regulated to Achieve Balance and Economy

    • Feedback inhibition: slows production by inhibiting catalytic activity when product accumulates

    • Systems biology: quantifies complex interactions among intermediates and pathways

  • Clicker Question 21: Feedback inhibition does what? A) contains the overall network. B) inhibits enzyme activity to slow product formation. C) is a sequence of consecutive reactions. D) increases production of intermediates.

  • Answer: B; feedback inhibition reduces enzyme activity to balance production

  • Clicker Question 21, Response: Inhibits catalytic activity to maintain balance in metabolic networks

  • Activity: Metabolic Map

    • further exploration of pathways and reactions in Achieve course

  • Activity Instructions (3 of 4)

    • address what drives electron flow to obtain energy; glycolysis energy source

  • Activity Solution (3 of 4)

    • redox reactions drive electron flow; ATP breakdown fuels endergonic steps

  • Bonus Discussion

    • identify intersections and shared intermediates; consider how changes in one pathway affect others


2. Chemical Foundations

  • Elements essential to animal life and health

  • Biomolecules are carbon-based with diverse functional groups

    • carbon forms single, double, and triple bonds; carbon skeletons define molecule properties

  • Geometry of carbon bonding

    • tetrahedral arrangement for four single bonds; free rotation around singles; limited rotation about double bonds

  • Common functional groups of biomolecules

  • Additional functional groups of biomolecules

  • Clicker Question 9: Important functional groups include: A) lipids. B) thioesters. C) nucleic acids. D) carbons.

  • Answer: B; thioesters are a notable functional group

  • Clicker Question 9, Response: Functional groups confer chemical properties; thioesters are one example

  • Polyfunctional biomolecules

  • Universal small molecules in cells

    • central metabolites: amino acids, nucleotides, sugars and phosphorylated derivatives, etc.

    • secondary metabolites: organism-specific

    • metabolome/metabolomics: study of small-molecule complement

  • Macromolecules are major cellular constituents

    • macromolecules = polymers > ~5,000 Da; major classes: proteins, nucleic acids, polysaccharides

    • oligomers = shorter polymers

    • informational macromolecules = proteins, nucleic acids, certain oligosaccharides with information-bearing sequences

  • Protein macromolecules

    • proteins = polymers of amino acids; functions include enzymes, structural roles, receptors, transporters

    • proteome/proteomics = protein complement & its study

  • Clicker Question 10: Proteins are macromolecules because: A) form large structures noncovalently. B) they are polymers with MW > ~5,000. C) they function as enzymes, structural elements, receptors, or transporters. D) they are composed of multiple oligomers.

  • Answer: B; MW > ~5,000 defines macromolecules

  • Clicker Question 10, Response: Correct

  • Nucleic acid macromolecules

    • DNA and RNA = polymers of nucleotides; store/transmit genetic information; some RNAs have structural/catalytic roles

    • genome = complete DNA/RNA sequence

    • genomics = genome structure/function/evolution/mapping

  • Polysaccharide macromolecules

    • polymers of simple sugars; roles include energy stores, cell-wall components, extracellular recognition

    • glycome = carbohydrate-containing molecules in a cell

  • Lipid molecules

    • lipids = hydrophobic hydrocarbons; roles include membranes, energy stores, pigments, signaling

    • lipidome = lipid-containing molecules in a cell

  • Activity: Classifying Metabolites

    • Palmitic acid structure exploration

  • Activity Instructions (2 of 4)

    • Determine if palmitic acid is carbon-based metabolite; what type of biomolecule is it?

  • Activity Solution (2 of 4)

    • Palmitic acid is carbon-based (16 carbons); it is a fatty acid, i.e., a lipid

  • Building Blocks of Biochemistry (reprise of major biomolecule classes)

  • Major Biomolecule Classes in E. coli (Table 1-1; percentages by weight)

    • Water ~70%

    • Proteins ~15% (~3,000 species)

    • Nucleic acids: DNA ~1%; RNA ~6% (~3,000+ species)

    • Polysaccharides ~3%; Lipids ~2%

    • Monomeric subunits and intermediates ~2%; Inorganic ions ~1%

    • Note: lipid diversity is enormous when considering fatty acid permutations

  • Clicker Question 11: The systematic characterization of the entire collection of small molecules in a cell under defined conditions is called: A) genomics. B) proteomics. C) lipidomics. D) metabolomics.

  • Answer: D; metabolomics

  • Clicker Question 11, Response: Metabolomics studies the metabolome under specified conditions

  • Three-Dimensional Structure: configuration and conformation

    • configuration = fixed spatial arrangement of atoms

    • stereoisomers = same bonds/formula, different arrangement

    • stereospecificity = interaction depends on conformation

  • Geometric isomers (cis/trans)

  • Chiral and Achiral molecules

    • chiral centers (asymmetric carbon)

    • a molecule with n chiral centers can have 2^n stereoisomers

  • Clicker Question 12: Molecules that differ in configuration cannot be: A) stereoisomers. B) cis-trans isomers. C) chiral centers that are mirror images. D) chiral centers that can be interchanged by rotation of a single bond.

  • Answer: D; rotation around a single bond can interconvert conformers, not configuration

  • Clicker Question 12, Response: Configuration vs. conformation distinction clarified

  • Enantiomers and diastereomers

    • enantiomers = mirror-image stereoisomers

    • diastereomers = non-mirror-image stereoisomers

  • Optical activity of enantiomers

    • enantiomers have similar reactivities but differ in optical rotation

    • racemic mixture shows no net optical rotation

  • RS naming for stereoisomers

    • priorities assigned to substituents; example ordering given in class notes for priority

  • Clicker Question 13 (summary): Enantiomers can exist for molecules with more than one chiral carbon

  • Answer: D; enantiomers exist when there are one or more chiral centers

  • Clicker Question 13, Response: D is correct; stereoisomerism arises from multiple chiral centers

  • Molecular Conformation

    • conformation = spatial arrangement of freely rotating substituents

  • Principle 2 (4 of 6) and related: re-emphasize the small set of metabolites and modular assembly of cellular structures

  • Interactions Between Biomolecules Are Stereospecific

  • Biological Systems Distinguish Stereoisomers

    • stereospecificity defined; biological recognition often enantioselective

  • 1.4 Genetic Foundations

  • Principle 4 (2 of 5): DNA stores information; precise replication and assembly guided by genome

    • bacterial replication can yield billions of identical daughter cells; vertebrate inheritance follows parental genes

  • Principle 2 (6 of 6) (reiteration for cross-reference)

  • Genetic Information Is Encoded in DNA

    • DNA = deoxyribonucleic acid; sequence of deoxyribonucleotides

    • encode instructions to form cellular components; provide a template to produce identical DNA

  • Genetic Continuity Is Vested in Single DNA Molecules

    • E. coli genome: single molecule ~4.64 million nucleotide pairs

    • replication must be near-perfect to yield identical progeny

  • The Structure of DNA Allows Replication and Repair with Near-Perfect Fidelity

    • deoxyribonucleotides = monomers of DNA

    • complementary base pairing across strands via hydrogen bonds

    • strands held together by hydrogen bonds; not covalent bonds between strands

  • Clicker Question 22: Which statement about DNA is false? A) DNA is a double helix of two polymeric strands. B) Each strand serves as a template for copying. C) Covalent interactions between complementary bases occur between strands. D) Deoxyguanylate is complementary to deoxycytidylate.

  • Answer: C; covalent interactions do not occur between complementary bases across strands; hydrogen bonds (noncovalent) do

  • Clicker Question 22, Response: The strands are held by hydrogen bonds (noncovalent); covalent bonds link bases to sugars within each strand

  • The Linear Sequence in DNA Encodes Proteins with 3D Structures

    • native conformation = precise 3D shape of a protein essential for function

  • Clicker Question 23: The DNA sequence of a gene for a protein:

    • A) is a nonlinear code. B) mutates at a high rate. C) ultimately determines the amino acid sequence of a protein. D) is a polymer of four ribonucleotides.

  • Answer: C; DNA sequence determines the amino acid sequence via transcription to RNA and translation

  • Clicker Question 23, Response: C is correct; the linear DNA sequence maps to a protein sequence which folds into a functional conformation

  • Activity: Relating Terms within the Chapter

    • concept map exercise to connect terms such as each cell, genomic information, precise self-replication, inheritance, etc.

  • Activity Instructions (4 of 4)

    • construct a concept map linking the listed terms

  • Activity Solution (4 of 4)

    • pair up to discuss variations and connections


3. Physical Foundations

  • Living organisms exist in a dynamic steady state, never at equilibrium; energy required to maintain order

  • Systems vs. Universe: open systems exchange both energy and matter with surroundings

  • Energy transformation in living systems

    • first law: energy conserves; energy changes form

    • second law: entropy tends to increase; local order maintained by energy input

  • Free energy and spontaneity

    • free energy change ∆G determines spontaneity; if ∆G < 0, reaction is spontaneous

    • relation: ∆G = ∆H − T∆S; where ∆H is enthalpy, ∆S is entropy

    • standard free energy change ∆G° and the actual ∆G depend on concentrations via: riangleG=riangleG+RTextlnQriangle G = riangle G^\circ + RT \, ext{ln} Q

  • Enzyme-catalyzed reactions

    • enzymes accelerate reactions by lowering activation energy (ΔG‡) without changing the overall equilibrium

    • reaction coordinate diagrams illustrate exergonic and endergonic steps and coupling

    • activation energy: riangleGriangle G^\ddagger

  • Coupling and Energy Transduction

    • ATP hydrolysis as a major exergonic driver for endergonic processes

    • energy coupling enables cellular work

  • Clicker Question 20 (review): Enzymes increase reaction rates but not equilibrium; answer B

  • Clicker Question 16 (review): Enthalpy change ∆H reflects bonds formed/broken

  • Clicker Question 15 (review): Open system for living organisms

  • Metabolic organization and regulation

    • metabolism = network of catabolic and anabolic pathways

    • flow of electrons (redox) powers energy capture and biosynthesis

    • catabolic → ATP, NAD(P)H production; anabolic uses energy for synthesis

  • Regulation via feedback inhibition

    • feedback inhibition maintains balance of intermediate production and utilization

    • systems biology approach to quantify interactions among pathways

  • Conceptual points for 1.3 Physical Foundations

    • reaction coupling is central to metabolism

    • free energy and disequilibrium enable biological work


4. Genetic Foundations

  • Principle 4 (2–5 of 5): Precise replication and assembly using genome information

    • bacteria: a single cell can give rise to many identical cells; vertebrates show resemblance across generations due to inheritance

  • Genetic Information Is Encoded in DNA

    • DNA = sequence of deoxyribonucleotides; code for forming cellular components

    • DNA provides a template to replicate identical DNA molecules

  • Genetic Continuity in Single DNA Molecules

    • E. coli genome size ~4.64 million base pairs in a single circular chromosome

    • faithful replication is essential for identical progeny

  • The Structure of DNA and Replication/Repair Fidelity

    • deoxyribonucleotides are the monomeric subunits; base pairing is complementary across strands via hydrogen bonds

    • strands are held together by hydrogen bonds; covalent bonds connect bases to the sugar-phosphate backbone within each strand

  • The Linear Sequence in DNA Encodes Proteins with 3D Structures

    • native conformation = precise 3D structure of a protein

    • DNA sequence encodes the amino acid sequence of a protein via RNA intermediate; protein folds into structure guided by sequence

  • Mutations and genetic variation

    • mutations alter nucleotide sequence and can affect cellular components; not all mutations are beneficial

  • The RNA role in prebiotic evolution (context)

    • RNA can act as catalyst and information repository; potentially first genes/catalysts

  • Miller–Urey experiments (context of origin of biomolecules)

    • simulated early-Earth conditions; energy sources included electrical sparks mimicking lightning

    • produced amino acids, hydroxy acids, aldehydes

    • false statement in quiz: A) energy provided by boiling the liquid phase is false; energy came from electrical sparks

  • The First Cell and Energy Sources (prebiotic evolution)

    • protocells formed from lipid vesicles containing self-replicating RNA

    • protocells with higher replication capacity became more numerous

  • Early energy sources for primitive cells

    • inorganic fuels (e.g., ferrous sulfide, ferrous carbonate)

    • photosynthesis later evolved; led to O2 in the atmosphere

  • Eukaryotic cells evolved from simpler precursors in stages

    • major changes: chromosome evolution, nucleus evolution, endosymbiotic events with aerobic/ photosynthetic bacteria

  • Endosymbiosis and mitochondria

    • eukaryotic mitochondria originated via symbiosis with ancient bacteria

    • mitochondria are integral to aerobic metabolism in modern eukaryotes

  • Evolutionary relationships revealed by molecular anatomy

    • homologs = proteins that share sequence similarity across organisms

    • sequence similarity informs phylogenetic relationships

  • Functional genomics and genomic organization

    • genes grouped by cellular processes; housekeeping genes expressed constitutively; regulatory genes modulate expression

    • differences in regulatory gene content contribute to organism complexity

  • Genomics and medicine

    • large-scale sequencing identifies genes linked to conditions; proteins encoded by these genes are drug targets

  • Clicker Question 25: Eukaryotic cells: A) first appeared about 3.5 billion years ago. B) obtained mitochondria via symbiotic bacteria. C) are similar to Archaeal cells in most respects. D) preceded the development of a double-membrane surrounding DNA.

  • Answer: B; mitochondria originated via endosymbiosis with an ancient bacterium

  • Clicker Question 25, Response: Endosymbiotic origin explains mitochondria presence in eukaryotes

  • Clicker Question 26: Genome sequencing showed that differences in human genes between individuals are relatively small; overall gene content is highly conserved across humans

  • Answer: D; sequencing shows few differences in genes between individuals, highlighting unity of life at molecular level

  • Activity: Evolution – Muddiest Point (1 of 2 and 2 of 2)

    • student activities to reflect on evolution concepts and clarify points of confusion

  • Summary points

    • molecular evolution connects genetic variation to phenotypic diversity

    • endosymbiosis is a key driver of eukaryotic complexity


5. Evolutionary Foundations

  • Principle 5 (2–3 of 3): Living organisms change over time by gradual evolution

    • vast diversity arises from shared ancestry; molecular evidence visible in gene/protein similarity

  • Changes in Hereditary Instructions Allow Evolution

    • mutation = changes in DNA sequence; can alter cellular components; may be beneficial or detrimental

    • wild type = unmutated baseline

  • Biomolecules First Arose by Chemical Evolution

    • Miller–Urey experiments showed plausible pathways to biomolecules from simple precursors under early-Earth conditions

  • The Role of RNA in Prebiotic Evolution

    • RNA could act as catalyst and information store; may have preceded DNA/proteins in early life

  • RNA or Precursors as First Genes and Catalysts

    • alternative scenarios include metabolic-first pathways or RNA-based systems

  • Miller–Urey experiments – false/true statements reviewed (context for exam)

    • false statement example: energy source not boiling; energy from electrical sparks

  • Biological Evolution Began > 3.5 Billion Years Ago

    • protocells with self-replicating RNA and lipid vesicles led to protocell lineages with replication advantage

  • First Energy Sources

    • early cells likely used inorganic fuels; photosynthesis evolved later to produce O2

  • Eukaryotic Cells Evolved via Endosymbiosis

    • mitochondria and chloroplasts trace to endosymbiotic events; nucleus evolution also pivotal

  • Molecular Anatomy Reveals Evolutionary Relationships

    • homologs across species indicate common ancestry; gene/protein sequence similarity reveals phylogeny

  • Genomic Comparisons in Medicine

    • genome sequencing identifies disease-associated genes; potential drug targets emerge from understanding gene function

  • Key Takeaways for Evolutionary Foundations

    • unity of life: despite diversity, core biomolecular machinery is conserved

    • evolution acts on variation via mutation and selection

    • endosymbiosis transformed prokaryotic cells into eukaryotes, enabling complex life

End of notes