Biotechnology
Journal
DOI 10.1002/biot.200600036 Biotechnol. J. 2006, 1, 822–827
822 © 2006 Wiley-VCH Verlag GmbH & Co. KGaA, Weinheim
1 Introduction
Oxygen is essential for maximal energy yield and optimal
utilization of substrate in every aerobic organism [1]. Dur-
ing growth of Aspergillus oryzae on solid substrates, the
aerial hyphae account for 70% of the oxygen uptake [2]. It
is shown that diffusion of oxygen is limited in the fila-
mentous fungal layer that covers the solid substrate and
that the substrate penetrative hyphae are limited in oxy-
gen consumption and growth [2, 3]. Therefore, oxygen
supply to filamentous fungal cells that are in close contact
with the substrate is considered as a bottleneck in solid-
state fermentation [2, 3].
Hemoglobins bind oxygen reversibly and have been
discovered in a wide range of organisms including verte-
brates, invertebrates, higher plants, fungi and bacteria
[4]. Despite the fact that all known hemoglobins have a
highly variable primary amino acid sequence, they all
show a six to eight α-helical arrangement that facilitates
binding of heme in the hydrophobic core of the protein [1].
Hemoglobin bridges a wide variation in oxygen tensions
at the sites of oxygen loading and unloading, and plays
therefore a major role in oxygen transport, although spe-
cific hemoglobins may be specialized for other particular
functions [4]. Several hemoglobin domain (HBD) genes
have been recently identified in various fungi, including
Aspergillus species (Te Biesebeke et al., An Aspergillus
oryzae gene expressed during hyphal growth is part of an
unusual family of fungal proteins, submitted). These HBD
Research Article
Expression of Aspergillus hemoglobin domain activities in
Aspergillus oryzae grown on solid substrates improves growth
rate and enzyme production
Rob te Biesebeke
1,2
, Amandine Boussier
1,2
, Nick van Biezen
2
, Machtelt Braaksma
2
,
Cees A. M. J. J. van den Hondel
3
, Willem M. de Vos
1,4
and Peter J. Punt
1,2
1
Wageningen Center for Food Sciences (WCFS), Wageningen, The Netherlands
2
TNO Quality of Life, Zeist, The Netherlands
3
Clusius Laboratory, Institute of Biology, Leiden University, Leiden, The Netherlands
4
Wageningen University and Research Center, Microbiology, Hesselink van Suchtelenweg, Wageningen, The Netherlands
DNA fragments coding for hemoglobin domains (HBD) were isolated from Aspergillus oryzae and
Aspergillus niger. The HBD activities were expressed in A. oryzae by introduction of HBD gene frag-
ments under the control of the promoter of the constitutively expressed gpdA gene. In the trans-
formants, oxygen uptake was significantly higher, and during growth on solid substrates the de-
veloped biomass was at least 1.3 times higher than that of the untransformed wild-type strain.
Growth rate of the HBD-activity-producing strains was also significantly higher compared to the
wild type. During growth on solid cereal substrates, the amylase and protease activities in the ex-
tracts of the HBD-activity-producing strains were 30–150% higher and glucoamylase activities
were at least 9 times higher compared to the wild-type strain. These results suggest that the As-
pergillus HBD-encoding gene can be used in a self-cloning strategy to improve biomass yield and
protein production of Aspergillus species.
Keywords: Aspergillus · Hemoglobin · Growth rate · Protein production · Solid-state fermentation
Correspondence: Dr.Rob te Biesebeke, Friesland Foods, P.O. Box 226,
8901 MA Leeuwarden, The Netherlands
E-mail: rob.tebiesebeke@frieslandfoods.com
Fax: +31-582992286
Abbreviations: CM, complete medium; DO, dissolved oxygen; HBD, hemo-
globin domains; PDA, Potato dextrose agar; WK, wheat kernels;
WSM, wheat-based solid medium
Received 15 March 2006
Revised 23 May 2006
Accepted 25 May 2006