GRADOS DE REDUCCION

Elemento Ƴ
C 4
H 1
N -3
O -2
P 5
S 6
GRADOS DE REDUCCION
SINTESIS CELULAR-RS (TEORICO)
dfefcd
 BIOENERGETICS OF MICROBIAL GROWTH 3

Table 2. Standard Gibbs Energy (G01

f
) and Enthalpy (Hf01 ) of Formation

Name Structure Phase G01

f kJ/mol Hf01 kJ/mol

H Proton H+1 aq −39.9 0.0

Hydrogen H2 g 0.0 0.0
O Oxygen O2 g 0.0 0.0
Water H2 O l −237.2 −285.8
N Dinitrogen N2 g 0.0 0.0
Ammonium NH+14 aq −79.4 −133.3
Nitrite NO−12 aq −32.2 −107.0
Nitrate NO−13 aq −111.3 −173.0
S Sulfur (rhombic) S s 0.0 0.0
Bisulfide HS−1 aq 12.1 −17.6
Thiosulfate S2 O−2
3 aq −522.5 −608.0
Sulfite SO−2
3 aq −486.6 −635.5
Sulfate SO−2
4 aq −744.6 −909.6
Fe Ferrous iron Fe+2 aq −78.9 −89.1
Ferric iron Fe+3 aq −4.6 −48.5
C Carbon monoxide CO g −137.2 −110.5
Carbon dioxide CO2 g −394.4 −393.5
Formate CHO−1 2 aq −351.0 −410.0
Bicarbonate CHO−1 3 aq −586.9 −692.0
Formaldehyde CH2 O aq −130.5 −150.2
Methyl amine CH3 NH+1 3 aq 20.7 −70.2
Methane CH4 g −50.7 −74.8
Methanol CH4 O aq −175.4 −245.9
Oxalate C2 O−2
4 aq −674.0 −
Glyoxylate C2 HO−1 3 aq −486.6 −824.0
Acetate C2 H3 O−1 2 aq −369.4 −485.8
Glycolate C2 H3 O−1 3 aq −517.6 −648.5
Acetaldehyde C2 H 4 O aq −139.1 −212.5
Glycinate C2 H4 O2 N−1 aq −324.3 −324.3
Ethane C2 H 6 g −32.9 −85.0
Ethanol C2 H 6 O aq −181.8 −288.3
Ethyleneglycol C2 H6 O2 aq −330.5 −
Malonate C3 H2 O−2 4 aq −700.0 −
Acrylate C3 H3 O−1 2 aq −286.2 −
Pyruvate C3 H3 O−1 3 aq −474.6 −596.0
Propionate C3 H5 O−1 2 aq −361.1 −510.4
Lactate C3 H5 O−1 3 aq −517.1 −687.0
b-Hydroxy propionate C3 H5 O−1 3 aq −518.4 –
Acetone C3 H6 O aq −159.8 −221.9
Alaninate C3 H6 O2 N−1 aq −509.9 −315.1
Hydrogen cysteinate C3 H6 O2 NS−1 aq −291.2 −349.6
Dihydroxy-acetone C3 H6 O3 aq −445.2 –
Propane C3 H8 g −24.0 −104.0
n-Propanol C3 H8 O aq −175.8 −331.0
2-Propanol C3 H8 O aq −185.3 −331.0
1,3-Propanediol C3 H8 O2 aq −327.0 –
Glycerol C3 H8 O3 aq −488.5 −676.0
Fumarate C4 H2 O−2 4 aq −604.2 −777.0
Oxaloacetate C4 H2 O−2 5 aq −793.8 −985.2
Sucinate C4 H4 O−2 4 aq −690.2 −909.0
Malate C4 H4 O−2 5 aq −845.0 −843.0
Tartrate C4 H4 O−2 6 aq −1010.0 –
Crotonate C4 H5 O−1 2 aq −277.4 –
Acetoacetate C4 H5 O−1 3 aq −493.7 –
Aspartate C4 H5 O4 N−2 aq −639.1 −906.4
Butyrate C4 H7 O−1 2 aq −352.6 −535.0
β-hydroxy butyrate C4 H7 O−1 3 aq −506.3 –
Glycylglycinate C4 H7 O3 N−1 aq −473.4 −690.3
Creatinine C4 H8 ON3 aq −23.2 –
Ethyl actetate C4 H8 O2 aq −355.0 −482.3
Acetoin C4 H8 O2 aq −280.0 –
4 BIOENERGETICS OF MICROBIAL GROWTH

Table 2. (Continued)

Name Structure Phase G01

f kJ/mol Hf01 kJ/mol

Creatine C4 H8 O2 N−13 aq −177.9 −128.2

Asparagine (dipolar) C4 H8 O3 N aq −526.3 −766.6
n-Butanol C4 H10 O aq −171.8 –
2,3-Butanediol C4 H10 O2 aq −322.0 –
Oxalosuccinate C5 H4 O−2
5 aq −1139.6 –
Glutamate C5 H7 O4 N−2 aq −643.9 −940.3
Valerate C5 H9 O−1
2 aq −344.3 −560.0
Valinate C5 H10 O2 N−1 aq −307.6 −567.8
Glutamine C5 H10 O3 N2 aq −528.4 −805.5
Citrate C6 H5 O−3
7 aq −1168.3 −1515.0
Cystinate C6 H10 O4 N2 S−2 aq −562.7 –
Caproate C6 H11 O−12 aq −336.0 –
Gluconate C6 H11 O−17 aq −1154.0 –
Leucinate C6 H12 O2 N−1 aq −296.8 −601.0
Glucose C6 H12 O6 aq −919.8 −1264.2
Mannnitol C6 H14 O6 aq −942.6 –
Benzoate C7 H5 O−1
2 aq −218.6 –
Sucrose C12 H22 O11 aq −1552.4 −2217.3
a-Lactose C12 H22 O11 aq −1565.9 −2233.8
a-Maltose C12 H22 O11 aq −1574.6 −2239.7
Biomass CH1.8 O0.5 N0.2 s −67.0 −91.0

formation values (Hf01 ) and Gibbs energy of formation similar, such that the organic fraction of biomass can be
values (G01 represented by a simple 1-C formula:
f ) as found in Table 2 as well as in Ref. 6.
The value for biomass is taken from (1). Although there
Biomass: C1 H1.8 O0.5 N0.2
is some discussion about the value of G01 f for biomass,
its exact value is not very important in thermodynamic This composition formula holds close to true for many
calculations, as shown by Heijnen et al. (7). organisms. When required one can determine the elemen-
The differences between microbial growth systems tal composition of the biomass and calculate a more precise
occur mostly in the electron acceptor/donor couples used. composition. For convenience we have left out the other
The most favorable inorganic N-source is NH+1 4 but some elements like P, S, K+ , and Mg2+ in our representation of
systems are capable of utilizing NO−1 −1
3 , NO2 , N2 or biomass, because of their minor contribution (<1–2%).
organic nitrogen as well. The biomass composition formula shows that synthesis
As shown in Fig. 1, the overall growth reaction can be of biomass requires a carbon source and a nitrogen source.
regarded as a combination of anabolism and catabolism. In addition, there is a need for an electron donor (D).
To establish the stoichiometry of the overall growth When growth is based on organic substrate degradation,
reaction, it is convenient to derive the stoichiometry of the then the organic substrate is both carbon and electron
catabolic and anabolic reaction first. In a second step in the donor (heterotrophic growth). When growth uses CO2 as
derivation of the overall growth reaction, the catabolic and C-source, then there is a need for a separate electron donor
anabolic reaction equations are coupled using one stoichio- to reduce CO2 to biomass (autotrophic growth):
metric coefficient, typically the biomass yield on substrate
(often equal to the carbon source and electron donor, • Heterotrophic:
ogr
1/YD ). In the following sections, we will first describe how
to derive the stoichiometry of the anabolic and catabolic a · electron donor/C − source + b · N
reaction. Then it will be shown how the overall growth − source + c · H+1 + d · H2 O + e · CO2
reaction can be derived as a resultant from the catabolism
+ 1 · CH1.8 O0.5 N0.2
and anabolism and one stoichiometric coefficient that is
typically based on bioenergetic considerations. • Autotrophic:

Stoichiometry of the Anabolic Reaction f · CO2 + a · electron donor + b · N − source

Microorganisms are composed of protein, Ribonucleic acid + c · H+1 + d · H2 O + e · oxidized electron donor
(RNA), Deoxyribonucleic acid (DNA), lipids, and carbo- + 1 · CH1.8 O0.5 N0.2
hydrates. Comparing many different (micro)organisms,
it is found that their relative contents are similar The unknown coefficients for the anabolic reaction
(40%–70% protein, 1.2% DNA, 5%–15% RNA, 2%–10% are easily calculated from the conservation requirements
lipid, 3%–10% carbohydrate). This similarity leads to (conservation of elements, electric charge), as shown in
an elemental composition of biomass which is also very Examples 1 and 2.