Chapter 21 : Glucose

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This chapter reviews the regulation of glucose uptake and metabolism in filamentous fungi and highlights the similarities with and differences from mechanisms in yeast. Fungal glucose transporters are classified as high-affinity transporters if the K for glucose is in the micromolar range and low-affinity transporters if K for glucose is in the millimolar range. contains a large number of proteins that can transport glucose across the yeast cell membrane, 17 of which (Hxt1 through Hxt11p, Hxt13 through Hxt17p, and Gal2p) belong to the yeast glucose transporter family. The use of glucose analogues has been used to determine whether glucose sensing in fungi requires uptake and/or further metabolism of glucose. The role of hexokinases in glucose sensing was confirmed in studies using strains carrying mutations in the three genes encoding sugar-phosphorylating enzymes: , , and . Given the central role of glucose in carbon metabolism and the diverse nutrient sources used by different fungi, it is not surprising that differences in glucose transport, glucose metabolism, glucose signaling, and carbon catabolite repression have arisen through selection.

Citation: Katz M, Kelly J. 2010. Glucose, p 291-311. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch21
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Image of FIGURE 1

Phylogenetic tree of putative sugar transporters in (AN) putative proteins that show similarity to fungal hexose transporters have been characterized in (Mst1), (HxtA and MstE), (MstA), (HGT1 and RCO3), (Hxt1p through Hxt11p, Hxt13p through Hxt17p, and Gal2p). (Gtt1), and (Hxt1p). The rooted tree was constructed using CLUSTAL ( ), PROTDIST and KITSCH ( ) through Biomanager at the Australian National Genome Information Service ((http://www.angis.org.au), and TREE-VIEW ( ). The amino acid sequences used to construct the tree were obtained from the Genome Database (http://www.yeastgenome.org/), NCBI (http://www.ncbi.nlm.nih.gov/), and the Comparative Database (http://www.broad.mit.edu/annotation/genome/aspergillus_group/MultiHome.html). The human glucose transporter GTR1 was included as an outgroup.

Citation: Katz M, Kelly J. 2010. Glucose, p 291-311. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch21
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Image of FIGURE 2

Enzymes of the glycolytic pathway and TCA cycle. In , genes encoding enzymes indicated in a solid box were increased, and those indicated in a dotted box decreased, in mycelia grown in glucose-rich compared to glucose-poor media ( ). Abbreviations: HEX, hexokinase/glucokinase; GPI, phosphoglucose isomerase; PFK, phosphofructokinase; FBA, fructose bisphosphatase; FBPA, fructose bisphosphate aldolase; TPI, triose phosphate isomerase; GPD, glyceraldehyde-3-phosphate dehydrogenase; PGK, phosphoglycerate kinase; PGM, phosphoglycerate mutase; ENO, enolase; PK, phosphoenolpyruvate kinase; PDC, pyruvate decarboxylase; ADH, alcohol dehydrogenase I; ALD, acetaldehyde dehydrogenase; ACS, acetyl-CoA synthase; PDH, pyruvate dehydrogenase; CT, citrate synthase; ACO, aconitase; IDH, isocitrate dehydrogenase; KDH, α-ketoglutarate dehydrogenase; SCoS, succinyl-CoA synthase; SDH, succinate dehydrogenase; FUM, fumarate dehydratase; MDH, malate dehydrogenase; PEPCK, phosphoenolpyruvate carboxykinase.

Citation: Katz M, Kelly J. 2010. Glucose, p 291-311. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch21
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Image of FIGURE 3

Phylogenetic tree of hexokinases and hexokinase-like proteins from , , , , , (formerly ) and . The rooted tree was constructed with amino acid sequences from the Fungal Genome Initiative at the Broad Institute (http://www.broad.mit.edu/annotation/fgi/) and sequences from ( ) by using CLUSTAL ( ), PROTDIST and KITSCH ( ) through Biomanager at the Australian National Genome Information Service (http://www.angis.org.au), and TREEVIEW ( ). Only two of the six sequences encoded by the genome were included ( ). Human glucokinase was included for comparison.

Citation: Katz M, Kelly J. 2010. Glucose, p 291-311. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch21
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Image of FIGURE 4

Conserved motifs within filamentous fungal carbon catabolite repressor proteins. Shown are sequence comparisons of ( [] EMBL ENCREA); ( EMBL AOR272151); ( EMBL ANCREA); ( EMBL AB024314); ( EMBL GFY16626); ( EMBL SSCRES); ( EMBL BCY16625); ( EMBL ACH245727); ( EMBL AB003106); ( EMBL TR27356); ( EMBL AF306571); ( EMBL MACRR1); ( EMBL AF055464); and ( EMBL THCRE1). Amino acid residues in , , and referred to in the text are underlined.

Citation: Katz M, Kelly J. 2010. Glucose, p 291-311. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch21
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Generic image for table

Characterization of glucose transporters in filamentous fungi

Citation: Katz M, Kelly J. 2010. Glucose, p 291-311. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch21
Generic image for table

Comparison of glucose sensing in and filamentous fungi

Citation: Katz M, Kelly J. 2010. Glucose, p 291-311. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch21
Generic image for table

Number of genes encoding hexokinases in filamentous fungi

Citation: Katz M, Kelly J. 2010. Glucose, p 291-311. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch21
Generic image for table

Comparison of mechanisms of regulation by major carbon catabolite repression proteins in fungi

Citation: Katz M, Kelly J. 2010. Glucose, p 291-311. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch21

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