Cloning and Expression of the XPR2 Gene from Yarrowia
lipolytica in Pichia pastoris
M. POZA, A. B. F. SESTELO, J. M. AGEITOS, J. A. VALLEJO,
P. VEIGA-CRESPO, AND T. G. VILLA*
Department of Microbiology, Faculty of Pharmacy, University of Santiago de Compostela,
15782 Santiago de Compostela, Spain
Yarrowia lipolytica is a dimorphic yeast able to secrete different types of proteases depending on the
pH of the environment. At neutral pH, the production of an extracellular alkaline protease (AEP) is
induced. This protease could be useful in the leather, detergent, or food industries. The XPR2 gene,
coding for AEP, was extracted from the pINA154 vector and cloned into the pHIL-D2 vector to obtain
a new protease-producing recombinant Pichia pastoris strain. The gene was efficiently integrated in
the P. pastoris genome and expressed from the AOX1 promoter actively induced by methanol. Finally,
the protease was successfully secreted by P. pastoris GS115.
KEYWORDS: Cloning; expression; protease; XPR2; Yarrowia lipolytica; Pichia pastoris
INTRODUCTION
Proteases are enzymes that catalyze the breakdown of the
peptide bonds in proteins and are involved in both normal and
abnormal biochemical pathways, conferring them the interesting
aspect of being potential targets for the development of new
therapeutic agents. Additionally, proteases are one of the three
main enzyme groups used industrially. Although these enzymes
are widely distributed in nature, in both animals and higher
plants, microbial proteases represent an interesting source with
clear industrial projection because of the rapid and profitable
growth of the microorganisms and their amenability to genetic
modification, this allowing the construction of new superpro-
ducing strains (1-3).
Yarrowia lipolytica is a dimorphic yeast able to secrete a
variety of enzymes including proteases. The type of protease
secreted depends on the pH of the environment. Thus, at acidic
pH, an acidic protease is produced (AXP), whereas neutral pH
induces the formation of an alkaline one (AEP) (4, 5). AEP
has been characterized as a serine protease, in particular, a
member of the subfamily of subtilisins with a molecular mass
of 30 kDa and an optimum pH of activity of 9-10 (6, 7). Its
corresponding gene (XPR2) has been cloned and sequenced,
resulting in a mature peptide encoded from an 890 bp sequence
preceded by an extensive pro-region including a signal peptide
(8). Alkaline proteases have been widely used for the detergents
industry, the best candidates being those who show broad
substrate specificity to facilitate the removal of a large variety
of proteinaceous substances. Also, both their stability at high
pH and their compatibility with other chelating and oxidizing
agents added to detergents are of great interest (3). Alkaline
proteases are also used in the leather industry, where extremely
alkaline conditions are often used for soaking, dehairing, or
bating. In addition, fungal proteases are often used in the food
industry to lower the bitter taste of protein hydrolyzates (3).
Pichia pastoris has routinely been used as an excellent host
for heterologous expression. In addition, the Pichia expression
systems allow the generation of stable recombinants microor-
ganisms where the DNA is permanently integrated in the
chromosomes (9), this being important for preventing gene
spreading (10). As an expression tool, P. pastoris has many
advantages, such as its economy of growth, the possibility of
making complex post-translational modifications, or the avail-
ability of a variety of plasmids as cloning vehicles (11, 12). In
addition, P. pastoris secretes low levels of native proteins, which
can ease the purification steps of the recombinant product. This
methylotrophic yeast is able to secrete larger than 30 kDa
heterologous proteins to a better extent than Saccharomyces
cereVisiae, where the cloned product is often hyperglycosylated
and hence may even lose its biological activity (5). The present
paper describes the cloning and expression in P. pastoris of an
alkaline protease (AEP) from Y. lipolytica.
MATERIALS AND METHODS
Strain, Plasmids, and Culture Media. The strains employed were
P. pastoris GS115 (his4 genotype, Pichia Expression Kit, Invitrogen,
Carlsbad, CA) and E. coli TOP10 (Zero Blunt Topo PCR Cloning Kit,
Invitrogen, Carlsbad, CA).
Plasmid pINA154 containing the XPR2 gene from Y. lipolytica was
obtained from Dr. Gaillardin (Collection de Levures d’Inte ´re ˆt Biotech-
nologique, UMR INA-PG-INRA-CNRS Microbiologie Ge ´ne ´tique Mo-
le ´culaire, INA-PG, Thiverval-Grignon, France) (7). Plasmid pCR-Blunt
II-TOPO, containing the kanamycin resistance gene (Invitrogen,
Carlsbad, CA), was employed to clone blunt-ends PCR products and
plasmid pHIL-D2 (Invitrogen, Carlsbad, CA) to express the XPR2 gene.
The pHIL-D2 vector is an integrative vector often used for intracellular
* Author to whom correspondence should be addressed [telephone +34
981592490; fax +34 981592490; e-mail mpvilla@usc.es].
3944 J. Agric. Food Chem. 2007, 55, 3944-3948
10.1021/jf0633894 CCC: $37.00 © 2007 American Chemical Society
Published on Web 04/14/2007