The Effects of Temperature on Life. Corien Bakermans, PhD Michigan State University
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1 The Effects of Temperature on Life Corien Bakermans, PhD Michigan State University
2 Overview Temperature Limitations Limits of Life Thermophiles and Psychrophiles Challenges and Solutions Nucleic Acids Proteins Cell Membranes Miscellaneous Summary
3 What is temperature? A measure of the average translational kinetic energy associated with the disordered microscopic motion of atoms and molecules.
4 Physical and Chemical Constraints High Temperatures: Fast, Fleeting, Flexible Reaction & diffusion rates increase Chemical degradation increases Flexibility/fluidity increase Liquid H 2 O Gas Low Temperatures: Slow, Stable, Solid Reaction & diffusion rates decrease Flexibility/fluidity of molecules decrease Liquid H 2 O Solid
5 Limits of Liquid H 2 O Increase boiling point: Increase pressure! Decrease freezing point: Add solutes!
6 Freezing Point Depression Seawater, 3% salinity: T m = -1.7 C Saturated NaCl brine: T m = - 21 C Other Brines Don Juan Pond CaCl 2 and NaCl brine T m = -52 C Mineral acids T m = -90 C As water solidifies, solutes are excluded from the crystal structure WEIGHT PERCENT NaCl
7 Known T Limits of Life Psychrophiles Hyperthermophiles Thermophiles Rothschild and Mancinelli (2001). "Life in extreme environments." Nature 409:
8 High T Limits of Life Enzymes active to 142 C Reproduction to 121 C strain 121, family Pyrodictiaceae (Kashefi and Lovley, 2003) Restricted to prokaryotic life! Bacteria max out at ~95 C Eucaryotes max out at ~60 C
9 Low T Limits of Life Reproduction to -12 C Eucaryotes -1.8C (freezing point of seawater) 200 Procaryotes -8 C Permafrost, Frozen Food isolates (Gilichinsky, 1995) C Psychrobacter cryohalolentis K5 (Bakermans, 2003) -12 C Psychromonas ingrahamii (Breezee, et al. 2004) 120 Metabolic activity to -20 C Eucaryotes 80 Himilayan Midge to -18 C (Kohshima, 1984) Antarctic lichens and endoliths (Friedmann, 1993) 40 Bacteria/Archaea Permafrost and South Pole snow (Rivkina, 2000; Carpenter, 2000) 0 Sea Ice (Junge, 2004) Can preserve cells at -196 C (liquid N 2 ) -40
10 Thermal Classes of Organisms
11 Temperature Effects on Growth Brock et al., 1994
12 High T Low T Nucleic Acids Proteins Lipids OTHER Denaturation by strand separation Degradation of bases (loss & deamination) Denature and aggregate metabolism collapses Deamidation of Asn & Gln Glu & Asp form succinimide His, Met, Cyst, Trp, & Tyr oxidized Too fluid Metabolites degrade Unfavorable secondary structures Difficult to unwind and access Cold denature (driven by hydration of polar and non-polar residues) Slowed reaction rates Too viscous, crystallize Ruptured by ice crystals Energy costs high
13 Nucleic Acids DNA RNA Ribosomal Messenger Transfer trna
14 Nucleic Acids at High Temperatures Problems: Strand separation Degradation of bases (loss and deamination) Solutions: Increase concentration of salts or polycationic polyamines (also help stabilize proteins) Introduce supercoils Cationic DNA-binding proteins Post-translational modification of trna DNA repair systems
15 Nucleic Acids at Low Temperatures Problems: Unfavorable secondary structures Difficult to unwind and access Solutions: DNA binding proteins RNA helicases to unwind secondary structure RNA chaparones Post-transcriptional modification of trna (increase dihydrouridine)
16 Proteins GOAL: balance molecular stability and structural flexibility in order to maintain activity
17 Protein Modifications Uncharged polar residues Charged residues Residue hydrophobicity Other Genome Level High T (thermophiles) sheets more likely residue volume size of proteins gln glu (H only) Low T (psychrophiles) (on surface) (neg. charges on surface) (but ) ion pairs & H bonds gly and pro gln thr leu??? hydrophobic res on surface??? Fields, 2001; Haney, PNAS, 1999; Chakravarty, 2000; Tekaia, 2002; Feller 1997; Saunders, 2003
18 Protein Stabilization Compatible solutes (aka extrinsic stabilizers ) Advantages: Can occupy broader thermal niches (simply vary concentration of solute) good for transient changes in T Affects all proteins in cell Examples: Increase with high temperature: 2,3-phosphoglycerate (at high T in methanogens) Myo-inositol phosphate derivatives, diglycerol phoshpate, mannosyl glycerate, inorganic ions, stress proteins Increase with low temperature: Stress proteins Glycine betaine, glycerol,trehalose, sucrose, proline, dimethylsuloniopropionate (Welsh, 2000)
19 Cell Membranes Fluidity is important for metabolic function. Fluidity affected by: Temperature Composition
20 Saturated Viscous Unsaturated Fluid
21 Maintaining Membrane Viscosity at High Temperatures (thermophiles) Archaea Ether bonds (vs. ester bonds) Enhance membrane packing to reduce fluidity: Caldarchaeols & nonitolcaldarchaeols Cyclic structures in transmembrane spanning fatty acids Bacteria Mimic archaeal ether & diether linkages with: Tetraesters Fatty alcohols Long-chain aliphatic diols Increase acyl-chain length, saturation, branching and/or cyclization
22 Membrane Lipids of Thermophilic Archaea
23 Maintaining Membrane Fluidity at Low Temperatures (psychrophiles) Reduce membrane packing to enhance fluidity Decrease acyl-chain length, saturation, and/or branching Saturated Unsaturated
24 Additional High Temperature Concerns Metabolites Problem: stability Very short half-lives at 95, 105 C FMN, pyridoxal phosphate, glucose-1,6-diphosphate, acetyl phosphate, CoASH, ATP, ADP Solutions: Protective microenvironment control ph, metal ion concentration, etc. Metabolic channeling between enzymes that are physically next to each other Alternate pathway or compound
25 Additional Low Temperature Concerns Ice crystals Live in a salty environment Produce cryoprotectants Antifreeze proteins Glycine betaine, glycerol, ectoine, proline, glutamate, alanine, trehalose (a.k.a. compatible solutes) Salt Produce compatible solutes Low diffusion rates More transporters in membranes Enzymes with higher affinity Energy More required at low T Energy Use (fmol/cell division) Energy Use Grow th Rate Temperature (Celsius) Growth rate (1/day)
26 Summary/Conclusions High Temperatures Fast, Fleeting, Flexible Low Temperatures Slow, Stable, Solid Have we identified the actual limits of terrestrial life and biomolecules? What factors will define the actual limits? High chemical degradation? Low presence of liquid H 2 0?
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