Urry D.W. (Ed.) What Sustains Life? : Consilient Mechanisms for Protein-Based Machines and Materials

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Results and Discussion

605

2.0

10^

•g

C/5

Kfi

i/j

1.5

1.0

5.0

xlO^

1.0x10^

10-

10^

Frequency (Hz)

10^ 10^

FIGURE 4. A. Discs of X2°-(GVGIP)32o. The upper

disc is impregnated with a propylene glycol and

water mixture and the lower disc contains water only.

Melding of Loss Shear Modulus and

Loss Permittivity Data to Evaluate

Acoustic Absorption Potential of

Elastic Protein-based Polymers

The temperature dependence of the loss shear

modulus data of Figure 4 is quite distinctive. At

200 Hz the temperature dependence is one of

increasing G' on increasing the temperature

from 15 to 25 C, but then the next larger G"

occurs at 40 C with the value of G' at

200

Hz for

the 30 C data being more than twice as large.

On the other hand at

MHz in Figure IB, the

increase in intensity of the loss permittivity

continues to increase on raising the tempera-

ture from 10 to

Nonetheless, the merging of loss shear

modulus, G", and loss permittivity, €\ data for

the overlapping frequencies in Figure 5 is truly

remarkable. Explanation of the distinctive 40 C

curve now becomes obvious. The explanation

arises due to the occurrence of the temperature

interval of the inverse temperature transition^

being from 10 to 30

During the inverse tem-

perature transition of hydrophobically folding

into the structure of Figure 2, the frequency

maximum of the loss permittivity peak

The lower disc is used in the collection of data in

figures 4B and 5. B. Loss shear modulus, G",

vs.

fre-

quency for (GVGIP)32o-

decreases as the elastic protein-based polymer

folds into a more regular structure. Once the

inverse temperature transition is complete, the

normal increase in frequency with increase in

temperature ensues.

A coarse calibration of the higher frequency

loss permittivity data becomes possible by using

the overlap with the loss shear modulus values.

It seems reasonable to conclude that the loss

shear modulus values would reach 5 x

10^

Pa or

more near

kHz. The comparison also suggests

that the intensity of the peak in Figure 1A would

almost double on continuing from 20 to 40

should also be appreciated that the composition

of the elastic protein-based polymer can be

chosen such that the temperature interval is

below 5 C in which case the intense absorption

would exist at temperatures above 5 C.

Mechanical Properties of

Cross-linked Fibers

When utilizing the apolar-polar repulsive free

energy for hydration, AGap, the free energy

gained on ion-pairing aligns and orders cross-

linking groups, E and K. This is demonstrated

for the following two pairs of polymers.

606

5.5

10^

Development of Elastic Protein-based Polymers as Materials for Acoustic Absorption

2.5 FIGURE

Melding of G" and e" data for

(GVGIP)320.

10"^

10"^ 10^ 10^ 10- 10^ 10^ 10^

I loss shear modulus | loss permittivity |

frequency (Hz)

Polymer

and for the

Polymer

x':

pair,

ii:

xi':

(GVGVP GVGVP GEGVP GVGVP GVGVP GVGVP)36 E/OF

(GVGVP)

(GVGVP GVGVP GKGVP GVGVP GVGVP GVGVP)22 K/OF

(GVGVP)

(GVGVP GVGFP GEGFP GVGVP GVGVP GVGVP)4o E/2F

(GVGVP)

(GVGVP GVGFP GKGFP GVGVP GVGVP GVGVP)22 K/2E

(GVGVP)

Micrographs of the fibers formed and repre-

sentative mechanical properties are given in

Figure 6. Among the several experiments,

values of the elastic modulus, E, at 20% of

greater than

10^

Pa have been obtained and

break stresses greater than 1.5 x

10^

Pa have

also been obtained for both fiber compositions.

These values are two orders of magnitude

greater than obtained by means or y irradiation

cross-linking. Generally, the more hydrophobic

phenylalanine (Phe, F) containing polymers

gave the higher values.

Essential Work of Fracture

The stress-strain curves given in terms of the

work in J/m^ versus extension to the point of

breakage is seen in Figure 7A. When the spe-

cific work of fracture is plotted as a function of

ligament length resulting from notching each

side of the elastic strip, extrapolation to the

intercept gives the essential work of fracture.

The results for both X'^-(GVGIP)26o and

X2o_(GVGIP)32o are shown in Figure

7B.

While

the gene for (GVGIP)26o indicates a chain

Results

and

Discussion

607

FIGURE

6. Mechanical properties

of chemically cross-linked fibers.

Polymer

ii~xi' Fiber

E (Pa) at 20%

Strain

3.86

10^

Break Stress

(Pa)

1.26

10'

Break

Strain

(%)

350

Stress-Strain Curves

Essential Work

Fracture (Intercept)

1400-

1400*

x2^-(GVGIP)

12 3 4 5

Ligament Length (mm)

FIGURE

Essential work

fracture

for

X2°-(GVGIP)26o

and

X2«-(GVGIP)32o.

608

Development of Elastic Protein-based Polymers as Materials for Acoustic Absorption

length of 260 pentamers, the expressed protein-

based polymer contained a smear of chains of

lower molecular weights. In making a new

monomer gene encoding for 10 pentamers, the

same codons were used for the two G residues

within the GVGIP pentamer but different

codons for G were used between the pen-

tamers. On multimerization a more stable

higher molecular weight gene product was

expressed, namely (GVGIP)32o. On 20 Mrad y-

irradiation cross-linking an elastic matrix

formed with twice the essential work of fracture

that was more consistent with chemically-syn-

thesized poly (GVGIP). Thus, a tougher matrix

was obtained that would be more suitable for

acoustic absorption applications.

Conclusions

Elastic protein-based polymers of the

(GXGXP)-type exhibit mechanical resonances

that open a new vista for the development of

unique acoustic absorption materials. Previ-

ously determined temperature dependent loss

factor (absorption/unit volume) curves for

X2'-(GVGIP)26o in the

100

Hz to

kHz range

have been dismissed as artifact in spite of the

fact that exactly parallel behavior as a function

of temperature is observed for a mechanical

resonance of (GVGIP)n at

MHz by means of

loss permittivity measurements. Melding of

loss shear modulus and loss permittivity

data confirmed the presence of an intense

mechanical resonance in the acoustic absorp-

tion range for matrices of X^^-(GVGIP)32o. The

cylindrical sample of Figure 8 stands ready for

acoustic absorption measurements by instru-

mentation claimed to be uniquely free of

artifact.

Using an understanding of the apolar-polar

repulsive free energy of hydration.

AGap,^

elastic

protein-based polymers can be prepared with

impressive mechanical properties of elastic

modulus and break stresses up to 2 x

10^

Pa and

break strains of several hundred percent.

Acknowledgments. The authors gratefully

acknowledge the Office of Naval Research

FIGURE 8. Cylinder of 20 Mrad y-irradiation cross-

linked (GVGIP)32o prepared in a form suitable for

acoustic absorption measurements by the "uniquely

artifact free" instrumentation.

under Contract Number N00014-00-C-0178 for

the support of these materials developments.

References

D.W.

Urry,

Hugel,

Seitz,

Gaub,

Sheiba,

J. Dea, J. Xu, and T. Parker, "Elastin: A Repre-

sentative Ideal Protein Elastomer," Phil. Trans.

Soc.

Lond.

B 357,169-184,2002.

R. Buchet, C.-H. Luan,

K.U.

Prasad, R.D. Harris,

and

D.W.

Urry,

"Dielectric Relaxation Studies on

Analogs of the Polypentapeptide of Elastin," /

Phys. Chem. 92, 511-517,1988.

D.W. Urry and T.M. Parker, MECHANICS OF

ELASTIN: Molecular Mechanism of Biological

Elasticity and its Relevance to Contraction, /

Muscle Res. Cell Motility 23, 541-547, 2002;

Special Issue: Mechanics of Elastic Biomole-

cules,

Henk Granzier, Miklos Kellermayer,

Wolf-

gang Linke, Eds.

CM. Venkatachalam and D.W. Urry, "Calcula-

tion of Dipole Moment Changes Due to Peptide

Librations in the Dynamic P-Spiral of the

References 609

Polypentapeptide of Elastin," Int. J. Quant

Chem.

Quant Biol Symp. 12,15-24,1986.

L. Onsager, Electric moments of molecules

in liquids. /. Am. Chem. Soc. 58, 1486-1493,

1936.

6. D.T. McPherson, J. Xu, and D.W. Urry, "Product

Purification by Reversible Phase Transition Fol-

lowing E. coli Expression of Genes Encoding up

to 251 Repeats of the Elastomeric Pentapeptide

GVGVP," Protein Expression and Purification 7,

51-57,1996.

D.W. Urry, K. Okamoto, R.D. Harris, C.E

Hendrix, and M.M. Long, "Synthetic, Cross-

Linked Polypentapeptide of Tropoelastin: An

Anisotropic, Fibrillar Elastomer," Biochemistry

15,4083^089,1976.

8. D.W. Urry, "Physical Chemistry of Biological

Free Energy Transduction as Demonstrated by

Elastic Protein-based Polymers," invited

FEATURE

ARTICLE, /. Phys. Chem. B, 101, 11007-11028,

1997.

9. B.E. Read, G.D. Dean, and J.C. Duncan, "Deter-

mination of dynamic moduli and loss factors."

In: Physical Methods of Chemistry, Vol. VII,

John Wiley & Sons, 1991.

10.

F. Prochazka, PhD Thesis: "Etude del'evolution

des relaxations dans les gels en cours de forma-

tion, appHcation aux polyurethanes" (1998) Uni-

versite du Maine, Le Mans—^France.

11.

T Nicolai, F. Prochazka, and D. Durand, "Com-

parison of Polymer Dynamics between Entan-

glements and Covalent Cross-links" Physical

Review Letters 82, 863-866,1999.

12.

M. Stading, M. Langton, and A.-M. Hermansson,

"Small and large deformation studies of protein

gels."/.

Rheol 39,1445-1450,1995.

Index

Abrahams, J.R, 405, 406, 408,409,

411,

416^18

Actin binding site

myosin cross-bridge use of,

440,

443-444

narrow cleft emanation and,

445^46

Adaptability, 28

Adenosine diphosphate

(ADP)

ATP-formed, 13, 320

repulsion burst and, 422

ATP produced by

phosphorylated, 46,

charged state bound, 4

phosphate energizing and

addition to, 349-350

phosphorylated, 46,47

Adenosine triphosphate (ATP),

346-351

ADP formed by, 13, 320

repulsion burst and, 422

ADP,

phosphorylated

production of, 46,47

analogue states, 446-447

apolar-polar repulsive free

energy bursts on

hydrolysis of, energy

conversions and, 546

binding, 315, 352-353, 425,

445^46,448, 518-519

conformational change and,

349

negative cooperativity and,

422^23

site,

314,445-446

clam-shaped globular protein

motif and cross-bridge

binding of, 57,58

cleft residence of, 57-59

contractile rotary motors

production of, 5

cross-bridge binding and,

57-58,433^39

cross-bridge state, detached of,

448

cusp of insolubihty, movable

and binding of, 315,

318-319

Dictostelium discoidium,

myosin motor domain, 12

elastic "pull" component of

force and apolar-polar

repulsion on binding of,

352-353

Fo-motor for synthase of, 51

Fi-motor for synthase of,

18-19,

51-52, 52

Fi-motor synthase of, "waters

ofThales" and, 19-20

folding/unfolding cycle

mediated, 308, 309

glucose oxidation for chemical

energy of, 68n27

hydrolytic rate, positive

cooperativity and, 423

hydrophobic domain repulsion

and, 311, 312-314

living organism construction/

maintenance and, 350-351

mechanical work and,

555-561

muscle relaxation and binding

of, 425

narrow cleft emanation and

binding site of, 445-446

P-,

y-phosphate of, 416^19

production, 4-5,46^7,

550-555

protein-based machines bound

to,

fully charged

carboxylate state and,

10-11

protein-based machines

energized by binding, 352

proton gradient and

production of, 550-555

push and pull production of,

4-5

repulsion, charged/

hydrocarbon group

produced by, 11

rigor mortis and, 245

rotary motor production of,

scallop muscle analogue states

of, 446-447

spHtting, 320

standard free energies and

phosphate bond

hydrolysis of, 347-349

P-subunit and y-rotor AG^p,

419^20

a-subunits

containing, 19,405,

410,411,415, 417

synthase, 19-20, 51-52

triphosphate tail binding of,

310

water availability to, 13

ADP,

Adenosine diphosphate

AFM. See Atomic force

microscope

611

612 Index

Alkalay, R.N., 504-506, 538

Allosteric effects, 21-22

AUosteric proteins

hydrophobic association

identification with

T-state

of, 243

R-state, 243

T-state, 243

Alzheimer's disease

amyloid deposits, 297

insolubiUty, excess and, 240

seminal amyloid-forming

sequence of, 303-304

solubility-insolubility

boundary, 3

Amino acid residues

aliphatic, slope source, 333-334

aromatic, slope source,

333-334

hydrophobic effect and, 333

hydrophobicity scale based on

T, for, 132-136

R-groups, 116,132,134-135

sequences, 125

control, 458

slope sources, 333-334

substitution, 132-133

Amino acids

hydrogen bond between

backbone peptide groups

and structure of, 125

hydrophobic association, 125,

126

residue protein return to free,

100

structures, 125

Trdivide, moving and

composition of, 115

valine replaced by, 115

Amyloid

deposits, 295

phenomenology of, 297

fiber formation, mechanism of,

297-304

formation, 295,297-304

peptide unit and prion protein,

299-300

prion protein, 298-300

Amyloidosis, 304

Amyloid precursor protein

(APP),

303-304

Analgesics, 511

Anemia, sickle cell, 279

Anesthetics, 511

Anfinsen cage function

mechanism, 316-318

Animal starch, 76

Animal waxes structure/

synthesis, 81

Antigen-antibody complexation,

131

Apolar-polar repulsion, 195-202

ATP binding, 352-353

hydrophobic face positioning

with, 415, 416

Apolar-polar repulsive free

energy, 339, 546

Apolar-polar repulsive free

energy of hydration. See

APP.

See Amyloid precursor

protein

Applied science, 456-457

Arrow of Time, 569

Arterial restoration, 23-24

Association (water insolubility),

Atomic force microscope (AFM)

bioelastic nanosensor for

kinase assays and, 565-566

single chain force extension

mode, 525-527

ATP.

See Adenosine triphosphate

ATPase motor, 53-56, 351

"idling," 13-14

ATPases, 13-14. See also Fr

motor ATPase

force generation in, 351-352

^Gap and, 351-353

myosin II motor, 351

skeletal muscle, 13

ATP synthase, 16, 394-424

breakdown, 394-395

c subunit of bovine, 402-404

electron density maps, 400-404

energy conversions, 394-395

Fi-motor

consilient mechanism

shown, 18-19

crystal structure, 404-420

Fi-motor component of, 17

function, 394-396

hydrophobic elastic consiUent

mechanism and, 423-424

protein subunits structural/

functional roles in,

398-399

structure

information, 400-420

schematic representation,

398^00

synthesis step, critical of, 424

thermodynamic data, 400-404

Back pain, lower, 465

Bacon, Francis (Sir), 547-548

The New Organon or True

Directions Concerning the

Interpretation of Nature,

571

Baromechanical transduction,

174

Beeswax synthesis, 81

Biocompatibility, 561-566

elasticity nature and,

130-131

elastic protein-based polymer,

561-562

protein-based polymer,

microbially produced,

489^95

resonances, medical and, 562

Biodegradable plastics, 25-26,

466-467, 533

biodegradation rates for,

25-26

model protein, 62, 66

thermoplastics, 25

Bioelastic disk, pressure ulcer

use,

523

Bioelastic matrices, 64-65

Biological interactions, 168

Biological structures

determination of, 76-90

proof,

synthesis, 76-90

Biology's Heat Engine, 542

Biology's machines, 50-59,

329-448.

See also specific

machines

Biology's Rival to Magnetism,

542

Biology's vital force, 541-542

elastic consilient mechanism

as mechanical coupler

within, 545-547

known, 561-566

physical embodiment of, 561

protein-based machines as

physical embodiments of,

548-561

Index 613

thermodynamics interlinked

with molecular dynamics

and, 545

Biomaterials, 561-566

protein-based polymer use as,

488-489

utility of advanced, 455^56

Biomolecular machines, 62-66

Biosensors, 525-533, 565-566

Biosynthesis, 561-566

protein-based polymer,

469^76

proteins of varied sequences

produced by, 30-32

Blood clotting

cusp of insolubility and,

283-294

fibrin clot, control of, 284-294

the Bohr Effect, 262-263

complex IV, 392

redox, 392

Boisvert, D.C., 311, 312, 327

Bovine spongiform

encephalopathy (BSE),

296

Boyer, P.B., 404, 412,453

Bronowski, J., 561

BSE. See Bovine spongiform

encephalopathy

Butler, J.A.V, 112,114,118,146,

147,172,181,185,201,

212,

213,264, 310, 325,

331,

334,425,449, 544, 572

CandeUUa wax synthesis, 81

Carbohydrates

animal starch, 76

cellulose, 76

D-glucose formation of, 76

photosynthesis production of,

354-355

plant starches, 76

structure, 76-77

synthesis, 76-77

Carbon (C), 29,72

mirror image molecules and

atom key of, 75-76

molecules containing, 74-75

Carboxylates, 115

Carboxylate state, fully charged,

10-11

Carboxyl ionization, 254-256

Carnauba wax synthesis, 81

Cell membranes

lipid bilayer assembly for, 78,

protein channels in, ion flow

through, 79-80

protein machine role in, 79-80

Cellulose, 76

Cerebral thrombosis, 3

CH2 units

solubility, water, 109-111

Trdivide moving by adding,

113-115

Chain molecules, 1

Chaos

energy inputs and, 228-235,

229-231

order created out of, 228-235,

566-568

reversing, 229-231

Chaperonins

consilient mechanism

exemplified by design of,

314-320

hydrophobicity unfolding/

refolding directed by,

308-314

molecular structure of, 305

substrates, 305-308

Charged groups (vinegar-like

groups)

ATP-produced repulsion

between hydrocarbon

groups and, 11

classic, 9

free energy and, 340, 341

hydrocarbon groups and, 11

competition for water

between,

6-10,18,123-124

hydrophobic hydration and,

119,

340, 341

Chemical energy, 32

acid, 221-222

ATP,

glucose oxidation for,

68n27

contraction driven by, 37

definition, 60-61

de-mixing achieved by, 221,

230-231

efficiency, 60-61

input

efficiencies, 206-208

temperature interval and

transition zone for,

152-154

light energy converted to, 44

mechanical work produced by,

151-156

motion produced by, 220-222

oil-like character of protein

increased by, 40

salt, 220-221

self-assembly achieved by, 221,

230-231

Chemical synthesis, 457-458,

476-482, 477^78

biocompatibility of protein-

based material, 478-482

DNA, 89-90

practicality, 478

protein-based material,

478-482

protein-based polymer,

476-482

repeating sequence protein-

based polymers solved

with, 473,477-478

Chemical work, 122

Chemochemical transducer,

398-400

Chemoelectrical transduction,

122

Chemomechanical transduction,

120,172

efficiencies, 342

efficiency statement for,

208-209

electrostatic charge-charge

repulsion and efficiencies

of, 208-210

myosin II motor, 555-561

Cholesterol synthesis, 80

Chronic wasting disease (CWD),

296

Cinnamide, 137

Citric acid cycle, 356

CJD.

See Creutzfeld-Jakob

disease

Clam-shaped globular protein

ATP analogue binding to

cross-bridge and, 57

contraction derived from

hydrophobically closed,

53,55

filament assembly, 52-53

muscle contraction as variation

on, 57, 58

scallop muscle and motif of,

56-59

614

Index

Cohen,

C, 448

Complex III, 330, 332, 336, 344,

359,

361-366, 368,

371-378,

386-392, 394,

439,452,

549-551,

561

Concatemerization,

471

Concatenation, monomer gene,

471

Consilient mechanisms, 26n5,

122-124, 212nl2,218

acceptance, 547-548

ATP synthase/Fi-ATPase

and

hydrophobic elastic,

423-424

biological, 329-448, 332-341

calcium iron trigger

and

hydrophobic, 446^48

chains, dynamic without

random chain networks

and elastic, 545-546

chaperonin design exempHfies,

314-320

chaperonin-directed

hydrophobic unfolding/

refolding and, 308-314

chemo-mechanical efficiencies

of electrostatic-induced

repulsion and, 209-210

definition,

329

denaturation, cold relationship

with,

202

de novo design origin

of, 2

diverse, 103-104

efficiency limit, 204-206

elastic, 331-332,423-424,455,

545-547

elastic hydrophobic, 423^24,

550

elasticity, energy conversion

and,127-128

electron transfer, 375-383

electron transport coupling

and,361-362

energy access

and

achieving

useful function

of,

219-224

of energy conservation,

131-147

energy conversions, 103-104,

332-341

energy resources

for,

131-147

function coupling

and,

166-167

AGHA,

169-170

physical basis

of,

176-195

energy resources, 137-146

Fo-motor

and

hydrophobic,

395

Fi-motor

and

hydrophobic,

395-398

Fi-motor synthesis

ATP

shown by, 18-19

function, 219-224

functional group coupling

and,

164-167

function coupling

and,

statement

energy

conversion, 166-167

genetic code relationship

to,

226-227

AGHA,

169-170

hemoglobin oxygen binding

and, 256-257

hemoglobin oxygen transport

and temperature effect

diagnostic

of, 254

hemoglobin sigmoid oxygen

binding and, 278

hydrophobic, 331-332,

395-398,

446^48,455,

545,548,

555-556

hydrophobic dissociation

relationship with,

202

hydrophobic elastic, 423-424,

550

inverse temperature transition

and hydrophobic, 545

mechanical work performed

by hydrophobic, 555-556

medical devices and,

562

myosin

motor, 430-446

oil-like domain association

controlled by, 103-104

Perutz mechanism melding

with, 278-279

protein-based machines

and

development

of,

547-548

diverse, 102-210

water required by, 11-13

as protein-based machines

biology, 329^48

protons transport,

at Qo and Qi

sites,

383

proton transfer, 375-383

push

and

pull,

4-5

Rayment

and

coworker

contributions to, 432^33

recognition,

soft tissue engineering,

562-563

temperature effect diagnostic,

282

as thesis

for

What Sustains

Life?,

2-A

"waters

Thales"

requirement

for, 336

Consihent protein-based

machines

efficiency, 203-210

energy conversions, 168-176

graphical representation, 35-37

motion produced

by,

158-160

Contractile machines.

See

also

Elastic-contractile model

protein machines

de novo-designed elastic model

proteins

of, 22

mechanical work performed

by,

nonmechanical work

performed by,

rotary,

Contractihty,

331

Contraction.

See

also Muscle

contraction

acid addition

and,

154-155

base addition

and,

155-156

chemical energy-driven,

clam-shaped globular protein,

hydrophobically closed

and deriving, 53,

elastic-contractile model

protein thermally driven,

111

elasticity

and,

128-130

electrical energy-driven,

energies driving,

32,

heat-driven, 154-158

ion pairing,

156

isometric,

344

mechanical work performed

by isometric,

344

model protein,

nicotinamide, lysine

attached reduction

driving,

222

and

energy required

drive,

reduction

and

heat-driven,

156-158

temperature increase

and

shortened,

Index

615

thermal energy-driven, 35-37,

120

transition zone for, 121-122,

154-156

Cooperativity, 308

ATP binding and negative,

422^23

ATP hydrolytic rate and

positive, 423

Harris and Rice treatment of,

200

hemoglobin positive, by

consilient mechanism, 278

hemoglobin subunit

interactions and positive,

253

Henderson-Hasselbalch

equation introduced into,

198-200

hydration competition and

positive, 338-339

hydrophobic domains and

positive, 254-257

hydrophobic interfaces and,

257,

260, 261

negative, 422^23

oxygen binding analogues to

formation of carboxylates

positive, 254-257

positive, 197, 253-257,

330-331,

338-339,423

Cortisone synthesis, 81

Creutzfeld-Jakob disease (CJD),

296,

297

Crick, F.H.C, 89

Crofts, A.R., 380,481

Cusp of insolubihty

anemia, sickle cell and lowered

temperature interval for,

279

blood clotting and, 283-294

definition, 2-3

excursions too far from, 3, 239,

241

Fi-motor, 18

hemoglobin transport of

oxygen and hydrophobic

association/dissociation

positioning of, 249-263

increased hydration around

oil-like groups lowers,

41-42

movement, 386

rotating, 18

temperature interval, 279

ubiquinone reduction and

movement of, 386

Cusp of insolubility, movable, 2,

3,18,241-263

ATP binding effect on, 315,

318-319

biology thrives near, 239-323

complex III and, 386

GroES capping effect on, 315,

318-319

hydration competition and, 18

molecular process visualized

by, 15-16

moving, 330

phosphate release and

temperature lowered for

substrate's, 320

representation of, 330

RIP movement and, 386

CWD.

See Chronic wasting

disease

Cytochrome bci complex, 359,

372,

448, 450,

451,

572

Cytochrome c oxidase, 386-394

electron transfer steps for

oxygen reduction with,

389

oxygen reduction with,

387-391

structure, 387

Cytosol, 383-384

Daniell, H., 464, 536

Dea,

214,

533,

537, 596, 599,

608

Denaturation

cold, 202

heat, 106,107

Deoxyhemoglobin, 253-254

crystal structure, 265-266

oxyhemoglobin external

differences from, 265-266

T state, 257

Deoxyribonucleic acid (DNA),

84.