Page 125 - Lignocellulosic Biomass to Liquid Biofuels
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Bioconversion of lignocellulosic biomass to bioethanol and biobutanol  99


              lignocellulose source. For instance, hemicellulose from a monocot source
              (grass) mainly includes xylose sugar: the furfural thus is the main furan
              generated in pretreated monocot; a significant amount of mannose is
              instead contained in the hemicellulose from softwood source: HMF thus
              is the most abundant furans in pretreated softwood lignocellulose [242].
                 According to research studies carried out by Aguilar [243] and Larsson
              [244], the furfural concentration in sugarcane bagasse can reach up to 5 g/
              L during acid hydrolysis [243], while the HMF can accumulate to 6 g/L
              in hydrolyzates from chipped pine wood [244]. Furfural and HMF are
              considered potent inhibitors of yeast cells growth and subsequent of fer-
              mentation process in a dose-dependent manner [245,246], causing the
              inhibition of sugar uptake rate and the decrease of ethanol production
              rate [247]. Anyway, they are usually toxic to many other types of organ-
              ism, such as mammalian cells, fungi, and bacteria [248]. These inhibitor
              compounds act by blocking both different enzymes of the glycolysis path-
              way, that is, hexokinase, phosphofructokinase, triose-phosphate dehydro-
              genase [239], and nonglycolytic enzymes. Modig et al. [248] reported the
              furfural and HMF inhibitory effects on three dehydrogenase enzymes:
              alcohol dehydrogenase (ADH), aldehyde dehydrogenase (AlDH), and the
              pyruvate dehydrogenase (PDH) complex. The relevant reactions carried
              out by these different enzymes are given by the following equations:
                                                  ADH
                                                          1
                                                1
                       Acetaldehyde 1 NADH 1 H  ! NAD 1 ethanol
                                          AlDH
                                         1
                      Acetaldehyde 1 NAD  ! acetic acid 1 NADH 1 H   1
                                        1  PDH                       1
               Pyruvic acid 1 CoA 1 NAD  ! acetyl-CoA 1 NADH 1 H 1 CO 2
                 Furfural and HMF can act synergistically, but the microbial activity of
              yeast strains seems to be more sensitive to inhibition by furfural than
              HMF, at the same concentration [245,249]. For instance, the presence of
              furfural negatively affects the activities of the ADH, AlDH, and PDH
              enzymes; in contrast, HMF has a less inhibitory effect on AlDH and PDH
              enzymes compared to furfural, while the effect on ADH enzyme is similar
              to that of furfural [248]. However, HMF can increase the lag phase of
              microbial growth, depleting the cell growth [250]. It was reported that in
              cultivating S. cerevisiae K35 and CBS 1200, and Z. mobilis ATCC 10988,
              5-HMF significantly inhibited cell growth and ethanol production [251].
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