BSC 450: Chapter 2

Water

2.1, Weak Interactions in Aqueous systems

Hydrogen bonding gives water its unusual properties
 Water has a higher melting point, boiling point, and heat of vaporization
than other solvents
 Attractions between adjacent water molecules gives liquid water great
internal cohesion
 Structure of the H2O molecule
o Each hydrogen atom shares an electron pair with the central oxygen
o Outer orbitals describe tetrahedron shape
o H-O-H bond angle is 104.5, slightly less than a normal tetrahedron
due to the crowding on nonbonding orbitals
o The oxygen nucleus is more electronegative than the hydrogen
 Results in two electric dipoles
 Each hydrogen bears a partial positive charge
 The oxygen bears a partial negative charge with 2x magnitude
 Hydrogen bond: an electrostatic attraction between the oxygen atom of one
water molecule and the hydrogen of another (three lines)
 Hydrogen bonds are relatively weak
o BDE = 23 kJ/mol (compared to covalent O-H bond with 470 kJ/mol)
o 10% covalent, 90% electrostatic
o Hydrogen bonds are constantly breaking and re-forming
 Tetrahedral arrangement allows one water molecule to form up to 4
hydrogen bonds
o Water: Due to continuous motion and disorganization, the avg. is 3.4
o Ice: each molecule is fixed in lattice structure and bonds with 4 atoms
 Hydrogen bonds account for the relatively high melting point of water
o Much thermal energy is required to break enough hydrogen bonds to
destabilize the crystal lattice
 When ice melts or water vaporizes, heat is taken up by the system
o The highly ordered arrays of water molecules in ice  less orderly
arrays in liquid water  wholly disordered gaseous state
o This occurs spontaneously, there is a push toward randomness (-ΔG)
 o ΔG represents the driving force
o ΔH is the enthalpy change from making and breaking bonds
o ΔS is the change in randomness

Water Forms Hydrogen Bonds with Polar Solutes

 Hydrogen bonds readily form between an electronegative atom and a
hydrogen atom covalently bonded to another electronegative atom
 Hydrogen acceptor: electronegative atom, usually N or O
 Hydrogen donor: electronegative atom covalently bound to H
 Hydrogens bonded to carbon atoms do not participate because carbon is only
slightly more electronegative than hydrogen (C-H bond is only weakly
polar)
 CH3CH2CH2CH3 has a BP of only 0.5 ° C
 CH3CH2CH2CH2OH has a BP of 117 ° C because it can form intermolecular
hydrogen bonds
 Uncharged polar biomolecules such as sugars dissolve readily in water
o Due to stabilizing effects of hydrogen bonds
o Alcohols, aldehydes, ketones, and N-H compounds all form hydrogen
bonds with water
 Hydrogen bonds are strongest when oriented to maximize electrostatic
interaction
o When the hydrogen atom and the two atoms that share it are in a
straight line (acceptor atom in line with covalent bond)
o Puts the positive charge on the H directly between the two partial
negative charges

Water Interacts Electrostatically with Charged solutes

 Water is a polar solvent
 Hydrophilic: compounds that easily dissolve in water
 Hydrophobic: nonpolar molecules
 Dissolving salts such as NaCl
o Water hydrates and stabilizes the ions, weakening electrostatic
interactions between them
 Water readily dissolves charged biomolecules
 Water replaces the solute-solute hydrogen bonds with solute-water hydrogen
bonds
 Water screens the electrostatic interactions between dissolved ions
o High dielectric constant
Entropy Increases as Crystalline Substances Dissolve
 As salts dissolve, ions leaving acquire greater freedom of motion
 Increase in entropy is responsible for the ease in dissolution
Nonpolar Gases are Poorly Water Soluble
 CO2, O2, and N2, are nonpolar
 Movement from gas  aqueous solution decreases entropy
 Water-soluble “carrier proteins”
o Facilitate transport of O2
o CO2 is transports as HCO3-
 NH3, NO, and H2S are polar and dissolve in water

Nonpolar Compounds Force Energetically Unfavorable Changed in the Structure

of Water
 All molecules dissolved in water interfere with hydrogen bonding of some
water molecules, but polar/charged solutes compensate by forming new
solute-water interactions
o Net change in enthalpy is generally small
 Hydrophobic compounds offer no compensation
o Small gain of enthalpy (requires input of energy from the system)
o Decrease in entropy (constrains water molecules in cage-like shells)
 Amphipathic compounds contain polar and nonpolar regions
o Polar region interacts with water and tends to dissolve
o Nonpolar regions avoid contact
o Hydrophobic effect: the nonpolar regions of amphipathic molecules
cluster to present the smallest hydrophobic area to the solvent, and the
polar regions are arranged to maximize their interaction with the
solvent.
o Micelles: stable structures of amphipathic molecules in water
o Hydrophobic interactions: the forces that hold the nonpolar regions of
the molecules together
 Result of minimizing the number of ordered water molecules
required to surround hydrophobic portions of the solute
molecules
o Aggregation of nonpolar amino acids in protein interiors stabilizes the
three-dimensional structure of proteins
o Release of ordered water favors formation of an enzyme-substrate
complex

Van der Waals Interactions Are Weak Interatomic Attractions

  When two uncharged atoms are brought close together, their electron clouds
influence each other
 Random variations in positions of electrons may create a transient electric
dipole, inducing a transient, opposite electric dipole in a nearby atom
 Van der Waals interactions: the weak attractions between two transient
dipoles, brings two nuclei closer together
 As the nuclei get closer together, their electron clouds repel each other
 Van der Waals radius: a measure of how close the atom will allow another to
approach

Weak Interactions Are Crucial to Macromolecular Structure and Function

 Van der Waals interactions, hydrophobic interactions, ionic interactions, and
hydrogen bonds are weak relative to covalent bonds but can have a
cumulative significant impact
 The formation in each weak bond contributes to a net decrease in the free
energy of the system
 To dissociate two noncovalently bonded molecules, all of the weak
interactions must be disrupted at the same time
o Constant random fluctuations make this very unlikely
 For macromolecules, the most stable structure is usually the one in which
weak interactions are maximized
o Folding polypeptide into three-dimensional shape
o binding antigen to antibody
o Enzyme binding to substrate
o Hormone biding to neurotransmitter
 For many proteins, tightly bound water molecules are essential to function
o Ex: cytochrome f has chain of five water molecules that provide a
path for protons to move across the membrane during photosynthesis
o Tightly bound water molecules can also be an essential part of a
protein’s ligand binding site

Solutes Affect the Colligative Properties of Aqueous Solutions

 Solutes can affect a solvent’s vapor pressure, boiling point, melting point,
and osmotic pressure (colligative properties)
 Due to the fact that concentration of water is lower in solutions than in pure
water
 Independent of the chemical properties of the solute, dependent on the
number of solute particles in a given amount of water
 o Ex: NaCl’s effect on osmotic pressure is twice that of a non-
dissociating solute such as glucose
 Water molecules tend to move from a region of higher water concentration
to one of lower water concentration (increase in disorder)
 When two solutions are separated by a semi-permeable membrane, water
molecules diffusing from a region of higher water concentration to lower
produces osmotic pressure
 Osmotic pressure: the force necessary to resist water movement
o pi=icRT
o R = gas constant, i= van’t Hoff factor (measure of the extent to which the solute
dissociates), ic= osmolarity
o Osmolarity: the product of the van’t hoff factor and the solute’s molar
concentration
 Ex: NaCl, i=2… glucose, i=1
o For solutions of several solutes, osmotic pressure is the sum of the contributiuons
of each species (pi=RT(i1ci + i2c2 + i3c3…)
 Osmosis: water movement across a semipermeable membrane driven by difference in
osmotic pressure
o Plasma membranes are more permeable to water than most other molecules and
ions due to protein channels (aquaporins)
o Isotonic: solutions of osmolarity equal to that of a cell’s cytosol
o Hypertonic: solutions of osmolarity greater than that of the cytosol
 She cell shrinks as water moves out
o Hypotonic: solutions of osmolarity lower than the cytosol
 The cell swells as water enters
o In their natural environment, cells generally contain higher concentrations of
biomolecules and ions than their surroundings
 Osmotic pressure tends to drive water into cells
 If not counterbalance, the inward movement of water will distend the
plasma membrane and cause bursting of the cell (lysis)
 In bacteria and plants, the cell wall prevents osmotic lysis
 Certain freshwater protists have a contractile vacuole that pumps
water out of the cell
 In multicellular atoms, blood plasma and interstitial fluid are
maintained at similar osmolarity to the cytosol (high concentration
of albumin and proteins in blood)
 Cells can actively pump ions such as Na+ into interstitial fluid to
stay in osmotic balance with surroundings
o Macromolecules has far less effect on osmolarity than would an equal mass of
their monomeric components
 Ex: 1g of polysaccharide composed of 1000 glucose units has the same
effect as 1mg of glucose
  Storing fuel as polysaccharides avoids an enormous increase in osmotic
pressure in the storage cell
o Plants use osmotic pressure to achieve mechanical rigidity
 High solute concentration in plant cell vacuole draws water in
 Nonexpendable cell wall prevents swelling
 Pressure exerted on cell wall (turgor pressure) increases
 Stiffens the cell, the tissue, and the plant body
 Loss of water reduces turgor pressure and causes the plant to wilt
o Laboratory protocols
 Mitochondria, chloroplasts, and lysosomes all have semi-permeable
membranes
 When isolating these organelles for research, you must work in isotonic
solutions to prevent entry of water, swelling, and bursting
 Buffers containing inert solutes such as sucrose protect organells against
osmotic lysis

2.2, Ionization of Water, Weak Acids, and Weak Bases

Pure Water is Slightly Ionized
 Water molecules have a slight tendency to undergo reversible ionization to
yield a hydrogen ion (proton) and hydroxide ion
o H2O  H+ + OH-
 Hydrogen ions formed in water are immediately hydrated to form
hydronium ions (H3O+)
 Hydrogen bonding between water molecules makes the hydration of
dissociating protons virtually instantaneous
 Ionization of water can be measured by its electrical conductivity
o Pure water carries electrical current as H3O+ toward the cathode, and
OH- toward the anode
o Extremely fast movement compared to other ions
o “proton hopping:” a series of protons hops between hydrogen-bonded
water molecules causes the net movement of a proton over a long
distance in a short time (OH- hops in the opposite direction)
 Acid-base reactions happen extremely fast
 Plays a role in biological proton-transfer reactions
 The position of equilibrium is given by its equilibrium constant, Keq
o A + B  C + D
o Keq = [C]eq [D]eq / [A]eq [B]eq
o The concentration terms should be the activities of each species, but
the equilibrium constant may be approximated by measuring the
concentrations at equilibrium
 o Unitless
o Fixed and characteristic for any given chemical reaction at a specific
temperature
 Defines final mixture regardless of starting reactants/products
 Can be calculated if concentrations of all reactants/products are
known

The Ionization of Water is Expressed by an Equillibrium Constant

 The degree of ionization for water at equilibrium is very small
 @ 25 C, only 2 of every 109 molecules ionize in any instant
 Kw is the ion product of water at 25 C
 Kw = 1.0 x 10-14
o So, the product [H+][OH-] will always equal Kw
 Kw = [H+][OH-]
 Neutral pH: when there are exactly equal concentrations of H+ and OH-
 When [H+] is very high [OH-] must be very low & vice versa

The pH Scale Designated the H+ and OH- concentrations

 pH= -log[H+]
 “p” means the “negative logarithm of”
 Acidic: pH less than 7
 Basic: pH greater than 7
 Neutral: pH precisely 7
 Logarithmic scale: difference of 1 pH unit is 10x the H+ concentration
 Indicator dyes
o Litmus, phenolphthalein, and phenol red
o Dyes undergo changes as protons dissociate from the dye molecule
 Electrodes
o Give accurate determinations of pH
o Sensitive to H+ concentration but insensitive to other cations
o Signal from glass electrode is amplified and compared with a solution
of known pH
 pH affects the structure and activity of macromolecules
o ex: catalytic activity of enzymes is strongly dependent on pH
o acidosis is when the blood plasma of people with sever diabetes falls
below a pH of 7.4
o alkalosis is when the pH of the blood is higher than normal

Weak Acids and Bases Have Characteristic Acid Dissociation Constants

  Strong acids and bases are completely ionized in dilute aqueous solutions
 Weak acids and bases are not completely ionized when dissolved in water
 Acids: proton donors
 Bases: proton acceptors
 Conjugate acid-base pairs
o When a proton donor loses a proton it becomes the corresponding
proton acceptor
 Each acid has a tendency to lose a proton in water, the stronger the acid, the
greater the tendency
o The tendency of HA to lose a proton and form A- is defined by the Keq
for the reversible reaction
 Keq = [H+][A-]/[HA] = Ka
o Equilibrium constants for ionization reactions are usually called
ionization constants or acid dissociation constants, Ka
o Stronger acids have larger ionization constants
o Weaker acids have smaller ionization constants
 pKa is analogous to pH
o pKa = log 1/Ka = -log Ka
o the stronger the tendency to dissociate a proton, the stronger is the
acid and the lower its pKa

Titration Curves Reveal the pKa of Weak Acids

 titration is used to determine the amount of an acid in a given solution
o A measure volume of acid is titrated with a solution of strong base of
known concentration
o The base is added in small increments until the acid is consumed
(neutralized)
 As determined with an indicator dye or pH meter
o The concentration of the acid can be calculated from the volume and
concentration of the base added
o Amounts of acid and base are often expressed in equivalents
 One equivalent is the amount of substance that will react with
or supply one more of hydrogen ions in an acid-base reaction
 Titration curve: a plot of pH against the amount of bas added
o Reveals the pKa of the weak acid
 Two reversible equilibria are involved in the process
o Before any base is added, the acid is already slightly ionized, to an
extent that can be calculated from its ionization constant
 o As NaOH is added, the added OH- combines with the free H+ in the
solution to from H2O to satisfy the equilibrium relationship
o As free H+ is removed, HA dissociates further to satisfy its own
equilibrium constant
o The net result as the titration proceeds it that more and more HA
dissociates, forming A-
o At the midpoint of the titration, at which exactly 0.5 equivalents of
NaOH have been added per equivalent of acid, one half of the acid has
undergone dissociation
 The concentration of the proton donor (HA) now equals the
proton acceptor (A-)
o At the midpoint: the pH of the equimolar solution of acid and
conjugate base is equal to the pKa of the acid
o As titration continues by adding more increments of NaOH, the
remaining non-dissociated acid converts into its conjugate base
o The endpoint of the titration all acid has lost its protons to OH- to
form H2O and its conjugate base
o Throughout the titration, the two equilibria coexist, each always
conforming to its equilibrium constant
 Higher Ka  stronger acid
 Lower pKa  stronger acid
 Stronger acid  loses proton more readily

2.3, Buffering Against pH Changes in Biological Systems

Buffers Are Mixtures of Weak Acids and Their Conjugate Bases

 Buffers: aqueous systems that tend to resist changes in pH when small
amounts of acid or base are added
 A buffer system consists of a weak acid (donor) and its conjugate base
(acceptor)
 Buffering region: the region on a titration curve where a flat zone of roughly
1 pH unit extends from the midpoint
o A given amount of H+ or OH- added has much less effect on pH in
here than it would outside of this zone
o At the midpoint, the concentration of the proton donor and the proton
acceptor are equal, maximizing buffering power of the system
 pH at midpoint is equal to pKa
 Strong acids and bases have no buffering power
  Buffering results from two reversible equilibria occurring in a solution of
nearly equal concentrations of a proton donor and conjugate proton acceptor
o When H+ or OH- is added to a buffer, the result is a small change in
the ratio of the relative concentrations of the weak acid and its anion,
causing a small shift in pH
o The decrease in concentration of one component is balanced by an
increase in the other
o The sum of the components does not change, only their ratio does
 Each conjugate acid-base pair has a characteristic pH zone in which it is an
effective buffer
o One pH unit above or below its pKa value

The Henderson-Hasselbalch Equation Relates pH, pKa, and Buffer Concentration

 The shape of the titration curve of any weak acid is described by the
Henderson-Hasselbalch equation
o pH = pKa + log [A-]/[HA]
 This equation shows why pKa of a weak acid is equal to the pH of the
solution at the midpoint of its titration, when [HA] = [A-]
o pH = pKa + log 1 = pKa + 0 = pKa
 This equation allows us to calculate:
o pKa, given pH and the molar ratio of proton donor/acceptor
o pH, given pKa and the molar ratio of proton donor and acceptor
o the molar ratio of proton donor and acceptor, given pH and pKa

Weak Acids or Bases Buffer Cells and Tissues against pH Changes

 Multicellular organisms have constant pHs
 Buffer systems are a first line of defense against changes to internal pH
 The cytoplasm of most cells contains high concentrations of proteins with
many amino acids with functional groups that are weak acids or weak bases
o Ex: histidine has a pKa of 6.0, so it can buffer effectively in neutral pH
(5-7)
o Ex: nucleotides such as ATP contain ionizable groups that contribute
to the buffering power of the cytoplasm
o Ex: organic acids buffer the vacuoles of plant cells
o Ex: ammonia buffers urine
 Two important biological buffers are phosphate and bicarbonate
o The phosphate buffer system:
 Acts in the cytoplasm of all cells
 H2PO4- is the proton donor & HPO42- is the proton acceptor
  H2PO4-  H+ + HPO42-
 Maximally effective close to its pKa of 6.86 (5.9-7.9)
 Most mammal’s cytoplasmic compartments have a pH in the
range of 6.9 to 7.4
o The bicarbonate buffer system:
 Buffers blood plasma
 HCO3- is the proton donor & HCO- is the proton acceptor
o H2CO3  H+ + HCO3-
o K1 = [H+][HCO3]/[H2CO3]
 This buffer system is more complex than other conjugate
acid-base pairs because one of its components, carbonic
acid (H2CO3) is formed from dissolved carbon dioxide
and water in a reversible reaction
o CO2(d) + H2O  H2CO3
o K2 = [H2CO3]/[CO2(d)][H2O]
 Carbon dioxide is a gas under normal conditions, and
CO2 dissolved in aqueous solution is in equilibrium with
CO2 in the gas (g) phase
o CO2 (g)  CO2(d)
o K3 = [CO2 (d)]/[CO2(g)]
 The pH of a bicarbonate buffer system depends on the
concentration of the proton donor and acceptor
components
 The concentration of H2CO3, the proton donor, depends
on the concentration of CO2 (d)
 The concentration of CO2 (d) depends on the
concentration of CO2 (g) or the partial pressure (pCO2)
 So.. the pH of a bicarbonate buffer exposed to a gas
phase is ultimately determined by the concentration of
HCO3- in the aqueous phase and by pCO2 in the gas phase
 The bicarbonate buffer system is an effective buffer near
pH=7.4
o Blood plasma is in equilibrium with a large reserve
capacity of CO2(g) in the air space of the lungs
o Reaction one and two occur in the aqueous phase
(blood in capillaries)
o Reaction three occurs going into the gas phase
(lung air space)
 Blood can pick up protons but also lose them
 Ex: gain of protons from lactic acid during exercise
 Ex: loss of protons by protonation of NH3 during protein
catabolism
 When H+ is added to the blood it passes through the tissues and
reaction 1 proceeds towards a new equilibrium
 H2CO3 increases, in turn increasing the CO2 (d) [reaction
2] in the blood, in turn increasing the pCO2 (g) in the air
space of the lungs [reaction 3], the extra CO2 is exhaled
 When H+ is lost from blood, the opposite occurs
 More H2CO3 dissociates into H+ and HCO3-, and thus
more CO2 (g) from the lungs dissolves in the blood
plasma
 The rate of respiration: the rate of inhaling and exhaling, can
quickly adjust these equilibria to keep the blood pH nearly
constant
 Controlled by the brain stem
 Detection of an increased blood pCO2 or pH triggers
deeper and more frequent breathing
 Hyperventilation: rapid breathing, tips the normal balance of O2
breathed in and CO2 out in favor of too much CO2 breathed out
 Raises the blood pH to 7.45 or higher
 This alkalosis can lead to dizziness, weakness, and
fainting
 Breathing into a paper bag
o Bag becomes enriched with CO2 and inhaling this
air increases the CO2 concentration in the body,
decreasing the blood pH
 At the normal pH of blood plasma (7.4), very little H2CO3 is
present relative to the HCO3-
 The addition of a small amount of base would titrate the
acid, exhausting its buffering capacity; this is
inconsistent with the large buffering ranges described
 Explanation for his paradox:
o The large reservoir of CO2(d) in blood, its rapid
equailibrium with H2CO3 results in the formation
of additional H2CO3
 In medicine it is useful to have a simple expression for blood
pH in terms of dissolved CO2
  Kh, the equilibrium constant for the hydration of CO2
 Kh = [H2CO3]/[CO2(d)]
 Ka = [H+][HCO3-]/[H2CO3] = [H+][HCO3-]/Kh[CO2(d)]
 KhKa = Kcombinesd = [H+][HCO3-]/[CO2(d)]
 Kh = 3.0 x 10-3
 Ka = 2.7 x 10-4
 Kcombined = 8.1 x 10-7M2
 pKcombined = 6.1
 blood pH = 6.1 + log20 = 7.4

Untreated Diabetes Produces Life-Threatening Acidosis

 Human blood plasma normally has a pH between 7.35 and 7.45
 Many enzymes in blood have evolved to have maximal activity in that range
 pH optimum: a characteristic pH where enzymes typically show maximal
catalytic activity
o Activity declines sharply on either side of this pH
o A small change in pH can make a large difference in the rate of some
crucial enzyme-catalyzed reactions
 Lack of insulin/insensitivity to insulin in people with untreated diabetes
disrupts the uptake of glucose from blood into the tissues an forces the
tissues to used stored fatty acids as fuel
o Results in accumulations of high concentrations of two carboxylic
acids (B-hydroxybutryric acid and acetoacetic acid)
o These acids have a combined blood level of 90 mg/100ml compared
to less than 3mg/100ml in healthy individuals
o These acids cause urinary excretion of 5g/24 hr compared with
<125/24 hr in control individuals
o Dissociation of these acids lowers the blood plasma pH to less than
7.35, causing acidosis and some enzymes to not function properly
 Diabetes mellitus is a likely diagnosis for a patient with high blood glucose,
low plasma pH, and high levels of the carboxylic acids in blood and urine
 Other conditions can produce acidosis:
o Fasting and starvation force the use of stored fatty acids with the same
consequences
o Heavy excretion in exercise leads to accumulation of lactic acid in
blood
o Kidney failure results in diminished capacity to regulate bicarbonate
levels
 o Lung diseases reduce the capacity to dispose of the CO2 produced by
fuel oxidation in the tissues, resulting in the accumulation of H2CO3
 Acidosis is treated by dealing with the underlying conditions

2.4, Water as a Reactant

Condensation Reactions
 The elements of water are eliminated
 Ex: formation of ATP from ADP and inorganic phosphate
Hydrolysis Reactions
 The reverse of condensation
 Cleavage accompanied by the addition of the elements of water
 Responsible for the enzymatic depolymerization of proteins, carbohydrates,
and nucleic acids
 Hydrolases: catalysis, invariably exergonic
o Produce two molecules from one, lead to an increase in the entropy of
the system
o Formation of polymers from subunits would be the reverse of
hydrolysis, but it would be endergonic and therefore doesn’t occur
 Cells can circumvent this obstacle by coupling endergonic
condensation reactions to exergonic processes, such as
breakage of the anhydride bond in ATP

Water in Reactions
 Water and carbon dioxide are the end products of the oxidation of fuels such
as glucose
o C6H12O6 + H2O  6CO2 + 6H2O
o The metabolic water formed is enough to allow some animals in very
dry habitats to survive
 The CO2 produced by glucose oxidation is converted in erythrocytes to the
more soluble HCO3-, in a reaction catalyzed by the enzyme carbonic
anhydrase
o CO2 + H2O  HC3- + H+
o Water is the substrate & functions in proton transfer my forming a
network of hydrogen-bonded water molecules through which proton
hopping occurs
 Green plants and algae use the energy of the sunlight to split water in the
process of photosynthesis
o 2H2O + 2A (light) O2 + 2AH2
 o A is an electron-accepting species, which varies with the type of
photosynthetic organism, and water serves as the electron donor in an
oxidation-reduction sequence that is fundamental to all life

2.5, The Fitness of the Aqueous Environment for Living Organisms

Organisms Have Adapted to Exploit the Properties of Water

 The high specific heat of water is useful to cells and organisms because is
allows water to act as a “heat buffer” keeping the temperature of an
organism relatively constant as the temperature of the surroundings
fluctuates and as heat is generated as a byproduct of metabolism
 The high heat of vaporization of water allows some vertebrates to lose
excess body heat to evaporate sweat
 The high degree of internal cohesion of liquid water due to hydrogen
bonding is exploited by plants as a means of transporting dissolved nutrients
form the roots to the leaves during transpiration
 The density of ice being lower than that of liquid water allows ponds to
freeze from the top down, the layer of ice on the top insulating the water
below from frigid air, preventing the pond from freezing solid
 Many physical and biological properties of cell macromolecules, particulary
the proteins and nucleic acids, derive from their interactions with water
molecules in the surrounding medium