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N Ahmed, S Shahab. Phosphate Solubilization: Their Mechanism Genetics And Application. The Internet Journal of
Microbiology. 2009 Volume 9 Number 1.
Abstract
The global necessity to increase agricultural production from a steadily decreasing and degrading land resource base has
placed considerable strain on agro ecosystems (Tilak, 2005). Current strategy is to maintain and improve agricultural
productivity exclusively via the use of chemical fertilizers. Although the use of chemical fertilizers is credited with nearly fifty
percent increase in agricultural production but they are closely associated with environmental pollution and health hazards
(Gaur and Gaind, 1999). Many synthetic fertilizers contain acids, such as sulfuric acid and hydrochloric acid, which tend to
increase the acidity of the soil, reduce the soil's beneficial organism population and interfere with plant growth. Generally,
healthy soil contains enough nitrogen-fixing bacteria to fix sufficient atmospheric nitrogen to supply the needs of growing plants.
However, continued use of chemical fertilizers may destroy these nitrogen-fixing bacteria. Furthermore, chemical fertilizers may
affect plant health. For example, citrus trees tend to yield fruits that are lower in vitamin C when treated with synthetic fertilizer.
Lack of trace elements in soil regularly dosed with chemical fertilizers is not uncommon. This lack of vital micronutrients can
generally be attributed to the use of chemical fertilizers. On the other hand Biofertilizer adds nutrients to soil.Environmentally
friendly biotechnological approaches offer alternatives to chemical fertilizers (Dobbelaere et al., 2003). Given the negative
environmental impacts of chemical fertilizers and their increasing costs, the use of PGPB is thus being considered as an
alternative or a supplemental way of reducing the use of chemicals in agriculture (De Weger et al., 1995; Gerhardson, 2002,
Postma, et al., 2003; Welbaum, 2004)
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Phosphate Solubilization: Their Mechanism Genetics And Application
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Phosphate Solubilization: Their Mechanism Genetics And Application
of radical orthophosphate from the carbon structure of the INORGANIC PHOSPHATE MINERALIZATION
molecule. The organophosphonates can equally suffer a Several reports have suggested the ability of different
process of mineralization when they are victims of bacterial species to solubilize insoluble inorganic phosphate
biodegradation (McGrath, 1995). The microbial compounds, such as tricalcium phosphate, dicalcium
mineralization of organic phosphorus is strongly influenced phosphate, hydroxyapatite, and rock phosphate (Goldstein,
by environmental parameters; in fact, moderate alkalinity 1986). In two thirds of all arable soils, the pH is above 7.0,
favors the mineralization of organic phosphorus (Paul and so that most mineral P is in the form of poorly soluble
Clark, 1988) The degradability of organic phosphorous calcium phosphates (CaPs). Microorganisms must assimilate
compounds depend mainly on the physicochemical and P via membrane transport, so dissolution of CaPs to Pi
biochemical properties of their molecules, e.g. nucleic acids, (H2PO4) is considered essential to the global P cycle.
phospholipids, and sugar phosphates are easily broken down, Evaluation of samples from soils throughout the world has
but phytic acid, polyphosphates, and phosphonates are shown that, in general, the direct oxidation pathway provides
decomposed more slowly (Ohtake, 1996; McGrath, 1995; the biochemical basis for highly efficacious phosphate
McGrath 1998). solubilization in Gram-negative bacteria via diffusion of the
strong organic acids produced in the periplasm into the
Phosphorus can be released from organic compounds in soil
adjacent environment. Therefore, the quinoprotein glucose
by three groups of enzymes:
dehydrogenase (PQQGDH) may play a key role in the
Nonspecific phosphatases, which perform dephosphorylation nutritional ecophysiology of soil bacteria. MPS bacteria may
of phospho-ester or phosphoanhydride bonds in organic be used for industrial bioprocessing of rock phosphate ore (a
matter substituted fluroapatite) or even for direct inoculation of
soils as a ‘biofertilizer’ analogous to nitrogen-fixing
Phytases, which specifically cause P release from phytic acid bacteria. Both the agronomic and ecological aspects of the
Phosphonatases and C–P Lyases, enzymes that perform C–P direct oxidation mediated MPS trait. (Gold stein et al., 2003)
cleavage in organophosphonates Among the bacterial genera with this capacity are
The main activity apparently corresponds to the work of acid Pseudomonas, Bacillus, Rhizobium, Burkholderia,
phosphatases and phytases because of the predominant Achromobacter, Agrobacterium, Microccocus, Aereobacter,
presence of their substrates in soil. Flavobacterium and Erwinia (Babu-khan et al 1995;
Goldstein, 1987; Sperber 1958; Rodríguez and Fraga, 1999).
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Phosphate Solubilization: Their Mechanism Genetics And Application
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Phosphate Solubilization: Their Mechanism Genetics And Application
solubilization in these strains is mediated by glucose or process of nitrate accumulation in soils might play an
gluconic acid metabolism. As solubilization of phosphate important role for the weathering of rock in general.
preceded detection of gluconic acid in the medium, perhaps
The nature and type of acid production is mainly dependent
even low levels of the acid (below the detection level of
on the carbon source (Reyes et al., 1999). In general, oxalic,
HPLC) started to dissolve the sparingly soluble phosphate.
citric, and gluconic acid, are strong solubilizing agents of
Alternatively, consumption of gluconic acid by growing
feldspar, biotite, and phyllosilicates, (Torre et al., 1993)
cells could also take place. In A. brasilense, reduction in the
quantity of soluble phosphate after incubation for 48 h can Figure 5
be explained as auto consumption of soluble phosphate by Table 1: Gluconic acid production by various bacterial
the growing bacterial population (Rodriguez et al., 2000). strains
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Phosphate Solubilization: Their Mechanism Genetics And Application
gibberllines and cytokinins (Kucey et al., 1988) are also Soils microorganisms are involved in a range of processes
correlated with Phosphate solubilization. Chelation involves that affect Phosphate transformation and thus influence the
the formation of two or more coordinate bonds between an subsequent availability of phosphate to plant roots
anionic or polar molecule and a cation, resulting in a ring (Richardson, 2001). Free living phosphate solubilizing
structure complex (Whitelaw, 2000). Organic acid anions, microorganisms (PSM) are always present in soils. The
with oxygen containing hydroxyl and carboxyl groups, have populations of inorganic Phosphate solubilizing
the ability to form stable complexes with cations such as microorganisms are sometimes very low, less than 102 CFU
Ca2+, Fe2+, Fe3+, and Al3+, that are often bound with phosphate -1
g of soil as observed in a soil in Northern Spain (Peix et al.,
in poorly forms (Jones ,1998 ; Kucey 1988) 2001). In four Quebec soils the number of root free PSM
-1
ranged from 2.5-to 3 x 106 CFU g of soil and they
Dissolution of phosphate in soil is a very important process
represented from 26- 46% of the total soil microflora
for plant growth. Several studies have shown that the
(Chabot et al., 1993). As observed with other soil microbes
phosphate uptake by plants can be markedly increased by
the number of PSM is more important in the rhizosphere
either mycorrhizal fungi (Azcon-Aguilar et al., 1986) or
than in non rhizosphere soil (Kucey et al., 1989), and the
inoculation of soil with species capable of solubilizing free
number of phosphate solubilizing bacteria is more important
phosphate, such as P. Bilaii (Cunningham and Kuiack 1992;
than that of fungi (Kucey, 1983). However, inoculation
Uzair et al., 2006).
studies aimed to improving P nutrition in plants involved
PHOSPHATE –PLANT INTERACTION bacteria and fungi, and is commercially available in Western
Canada as the phosphate inoculant Jumpstart (Philom Bios,
Phosphorus is one of one of the major plant nutrient limiting
Saskatoon, Sask.). They are sold for wheat, canola, mustard
plant growth. It plays a key role in nutrition of plants as it
and other legumes and contain an bacterial strain of
promotes development of deeper roots. The average soil is
Penicillium Bilaii. (http://www.philombios.ca/).
rich in phosphorus as it contains about 0.05% (w/w)
phosphorus (Barber, 1984) but only one tenth of this is PLANT GROWTH PROMOTING BACTERIA
available to plants approximately 95–99% is present in the
Although plant growth promoting bacteria occur in soil,
form of insoluble phosphates and hence cannot be utilized
usually their numbers are not high enough to compete with
by the plants and due to its poor solubility and chemical
other bacteria commonly established in the rhizosphere.
fixation in the soil (Gaurand Gaind, 1999) causing a low
Therefore, for agronomic utility, inoculation of plants by
efficiency of soluble P fertilizers.
target microorganisms at a much higher concentration than
To increase the availability of phosphorus for plants, large those normally found in soil is necessary to take advantage
amounts of fertilizer is used on a regular basis. But after of their beneficial properties for plant yield enhancement.
application, a large proportion of fertilizer phosphorus is (Igual, 2001)
quickly transferred to the insoluble forms. Therefore, very
little percentage of the applied phosphorus is used, making
continuous application necessary. (Abd Alla, 1994).
Figure 6
Figure 5: Soil and root interactions
(http://www.sare.org/publications/bsbc/fig3_3.jpg)
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Phosphate Solubilization: Their Mechanism Genetics And Application
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Phosphate Solubilization: Their Mechanism Genetics And Application
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Phosphate Solubilization: Their Mechanism Genetics And Application
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Phosphate Solubilization: Their Mechanism Genetics And Application
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Phosphate Solubilization: Their Mechanism Genetics And Application
in AG121 (Babu khan et al., 1995: krishnaraj, 2001). These dehydrogenase. PQQ belongs to the family of quinone
clones have no sequence homology to known PQQ genes but cofactors that has been recognized as the third class of redox
at least two have some homology to trans-membrane or cofactors following pyridine nucleotide- and flavin-
periplasmic proteins that could mediate an environmental dependent cofactors.
signal. Goldstein develops a strategy to open the funnel,
PQQ is a prosthetic group required by several bacterial
thereby increasing the rate of catalysis with a concomitant
dehydrogenases, including methanol dehydrogenase (MDH)
increase in gluconic acid production and CaP solubilization.
of Gram negative methylotrophs and some glucose
(Goldstein, 2003)
dehydrogenases. PQQ is derived from two amino acids,
GENETIC BASIS OF INORGANIC PHOSPHATE tyrosine and glutamic acid (Houck, 1991,Van Kleef, 1988),
SOLUBILIZATION but the pathway for its biosynthesis is unknown.
The genetic basis of mineral phosphate solubilization i.e. the
MPS phenotype is not well understood (Goldstein and Liu, Figure 13
1987). Because the production of organic acids is considered Figure 10: Alignment of pqq gene cluster. Equivalent genes
to be the principal mechanism for mineral phosphate have the same pattern
solubilization, it could be assumed that any gene involved in
organic acid synthesis might have an effect on this character.
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Phosphate Solubilization: Their Mechanism Genetics And Application
identified, indicating that PQQ may function as a vitamin in It has been concluded that organisms like E. coli and S.
mammals as well (Duine1999). typhimurium lack the genes to synthesize PQQ and it
remains unclear what the role of such apo-GCDs could be. It
PRESENT STATUS, DISTRIBUTION AND has been suggested that organisms that are unable to
SIGNIFICANCE OF PQQ
synthesize PQQ could scavenge it from their surroundings
PQQ was discovered in 1979 from a bacterium, and and activate their apo-quinoproteins (Matsushita & Adachi,
afterward it was reported to be in common foods. Because 1993).
PQQ-deprived mice showed several abnormalities, such as
poor development and breakable skin, PQQ has been BIOSYNTHETIC ROUTE OF PQQ
considered as a candidate for vitamin. It was a mystery, that The biosynthetic route of PQQ has not been elucidated yet,
until 2003 it was not identified as vitamin. Since the first but it has been proposed that glutamate and tyrosine are
vitamin (now called vitamin B1) was discovered in 1910 by precursors of PQQ (Houck et al., 1988, 1991; Van Kleef &
Dr. U. Suzuki, thirteen substances have been recognized as Duine, 1988). In all cases studied, the pqq operon contains a
vitamins; the latest one was vitamin B12 found in 1948.So it small gene; in the case of K. pneumoniae, this is pqqA
takes 55 years to discover “PQQ” a previously identified which can encode a small peptide of 23-29 a containing a
substance as new vitamin ( Choi 2008; Kashara and glutamate and tyrosine residue at conserved positions.
kato2003). Synthesis of the small peptide has been demonstrated
(Velterop, 1995) and it is thought to be the precursor of PQQ
After it had been established that PQQ occurs in several
(Goosen et al., 1989, 1992; Meulenberg et al., 1992). The K.
bacterial enzymes, a logical next question was whether it
pneumoniae pqqF gene product shows similarities to
also occurs in higher organisms. Perhaps stimulated by the
proteases like E. coli protease III (or pitrilysin) and
reports (Paz, 1988) that PQQ occurs at high levels in certain
insulinases and it has been proposed that the PqqF enzyme is
body fluids and tissues of mammalian milk, and in citrus
involved in cleavage of the PqqA precursor (Meulenberg et
fruits, several reports followed in which beneficial effects
al., 1992).
were ascribed to its administration, e.g. that a diet
supplemented with PQQ improved the “health” of mice Expression of the six K. pneumoniae pqq genes in E. coli
substantially or prevented the outbreak of certain diseases results in PQQ synthesis and an active GCD (Meulenberg et
(Duine, 1999). al., 1990, 1992). Most likely PQQ biosynthesis takes place
in the cytoplasm and PQQ can be released into the medium.
The quinoprotein glucose dehydrogenase (GCD) has been
Since the PQQ binding site of GCD faces the periplasm
demonstrated in a number of microorganisms including the
(Yamada et al., 1993), it is clear that both complementation
enteric bacteria Klebsiella pneumoniae, E. coli and
of E. coli with the K. pneumoniae pqq genes as well as
Salmonella typhimurium, and in Acinetobacter,
addition of PQQ can result in an active GCD (Matsushita,
Pseudomonas, Agrobacterium and Gluconobacter species. In
1997). From growth studies with E. coli ptsHI mutants
most cases, GCD is a membrane bound enzyme which
complemented with the complete set of K. pneumoniae pqq
oxidizes glucose to gluconate in the periplasmic space
genes or with plasmids lacking one of the six pqq genes and
(Duine, 1991). The formed gluconate is subsequently taken
from measurement of PQQ biosynthesis in the growth
up in a number of organisms via a specific transport system
medium, it was concluded that each of the six genes is
and further metabolized. The reducing equivalents from
required for PQQ biosynthesis (Velterop et al., 1995).
glucose are donated via PQQ to the respiratory chain.
Surprisingly, in some of the organisms mentioned above In Methylobacterium extorquens AM1, the genes for PQQ
only apo-GCD is found. Thus, K. pneumoniae synthesizes a synthesis are found in two clusters, pqqAB (C/D) E and
holo-GCD (Neijssel et al., 1983) but the closely related E. pqqFG (Toyama et al., 1997). These gene designations
coli and S. typhimurium can synthesize only apo-GCD but standardize the nomenclature with that of K. pneumoniae.
not PQQ (Hommes et al., 1984, 1986). However, apo-GCDs These genes in Methylobacterium strains were formerly
in these and other organisms can be converted into active called pqqDGCBA (Morris et al., 1994) and pqqEF
holo-GCDs by the addition of extracellular PQQ (Ameyama (Springer et al., 1996), respectively. In M. extorquens AM1,
et al., 1986; Hommes et al., 1986, 1984; Van Schie et al., pqqC and pqqD are not separate genes. Instead, they are
1987). fused into a single gene, pqqCD (Toyama et al., 1997).
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Phosphate Solubilization: Their Mechanism Genetics And Application
Transcriptional analysis of pqqA in M. extorquens AM1 peptide of 22-29 amino acids, which contains conserved
showed that a short mRNA containing only pqqA was tyrosine and glutamate residues. Since tyrosine and
produced at a much higher level than the mRNA for pqqAB glutamate are the probable precursors for PQQ synthesis
(Ramamoorthi & Mary, 1995). Similar results were obtained (Van Kleef and Duine, 1988; Houck et al., 1991), it has been
in the case of K. pneumoniae by using pqqTn5::lacZ operon proposed that this peptide is the precursor from which PQQ
fusions on the chromosome (Velterop et al., 1995).These is synthesized (Goosen et al., 1992). However, the
results are also consistent with the hypothesis that pqqA is a biochemical steps of PQQ synthesis are still unknown.
PQQ precursor, because stoichiometric amounts rather than Velterop et al., (1995) examined PQQ synthesis in vitro.
catalytic amounts are required if the peptide is the substrate
A series of experiments was carried out in which cell
for PQQ synthesis (Velterop et al .,1995)..
extracts of Escherichia coli containing all but one of the Pqq
GENETICS OF PQQ proteins were combined with those containing the missing
Genes involved in PQQ synthesis have been cloned from Pqq protein. PQQ was produced in only one of these sets,
Acinetobacter calcoaceticus (Goosen et al., 1989), K. that involving PqqC. E. coli cells containing a clone
pneumoniae (Meulenberg et al., 1992), Pseudomonas encoding all but the PqqC protein apparently produced an
fluorescens CHA0 (Schnider et al., 1995), Methylobacterium intermediate of PQQ, found both in the culture medium and
organophilum DSM 760 (Biville et al., 1989) and M. in the cells. However, the mount of the intermediate was low
extorquens AM1 (Morris et al., 1994). In A. calcoaceticus, and it was unstable (Velterop1995).
five pqq genes were identified and sequenced, designated
IV, V, I, II and III (Goosen et al., 1989). In K. pneumoniae, {image:14}
genes analogous to those were identified and designated
pqqABCDE, and in addition, a sixth gene was found (Source: Felder et al., 2000)
immediately downstream of pqqE, designated pqqF Genes involved in PQQ biosynthesis have been cloned from
(Meulenberg et al., 1992) In Methylobacterium strains, a several organisms. Five A. calcoaceticus pqq genes, pqqIV,
five gene cluster (designated pqqDGCBA) was identified by V, I, II, and III (Goosen et al., 1989, Goosen et al., 1987),
complementation analysis (Biville et al., 1989; Morris et al., and six K. pneumoniae pqq genes, pqqA, B, C, D, E, and F
1994), and sequence data from M. extorquens AM1 showed (Mellenberg 1990, Mellenberg et al., 1992), were cloned and
that the first three of these genes (pqqDGC) were analogous sequenced. Comparison of the deduced amino acid
to pqqABC of K. pneumoniae (Morris et al., 1994). In P. sequences showed that the proteins encoded by the first five
fluorescens, genes analogous to pqqFAB of K. pneumoniae genes of the K. pneumoniae pqq operon (pqqABCDE) show
have also been sequenced (Schnider et al., 1995). similarity to the proteins encoded by the corresponding A.
calcoaceticus genes (49 to 64% identical amino acid
Figure 14 residues). The K. pneumoniae pqqF gene encodes a protein
Table 3: Percentage similarities between pqqBCDE genes of that shows similarity to E. coli protease III and other
different bacteria proteases (Mellenberg et al., 1992), but its equivalent has not
yet been found in A. calcoaceticus. Recently, three M.
extorquens AM1 pqq genes, pqqD, G, and C, have been
cloned and sequenced (Morris et al., 1994); pqqC was only
partly sequenced. The encoded proteins showed similarity to
the K. pneumoniae PqqA, B, and C proteins and the A.
calcoaceticus PqqIV, V, and I proteins, respectively. Four
additional pqq genes have been detected in M. extorquens by
isolation of mutants and complementation studies. From
similar studies, six (possibly seven) pqq genes have been
postulated in M. organophilum DSM760 (Biville, 1989).
Finally, a DNA fragment cloned from Erwinia herbicola
(Source: Felder et al., 2000)
contained a gene encoding a protein similar to K.
In all four sequences, a small gene is present that encodes a pneumoniae PqqE and A. calcoaceticus PqqIII (Liu, et al.,
13 of 19
Phosphate Solubilization: Their Mechanism Genetics And Application
1992). Except for the K. pneumoniae PqqF protein, none of on yield of sorghum [Sorghum bicolor (L.) Moench] in dry
land. Trop.Agric. 69, 347–350
the Pqq proteins shows similarity to other proteins in the r-3. Ameyama, M., M. Nonobe, E. Shinagawa, K.
database. One of the pqq genes is small and may encode a Matsushita, K. Takimoto and O. Adachi, 1986. Purification
polypeptide of 24 amino acids (PqqIV, A. calcoaceticus), 23 and characterization of the quinoprotein d-glucose
dehydrogenase apoenzyme from Escherichia coli. Agric Biol
amino acids (PqqA, K. pneumoniae), or 29 amino acids Chem 50: 49-57.
(PqqD, M. extorquens AM1). Interestingly, these putative r-4. Antoun, H. and J.W. Kloepper, 2001. Plant Growth
promoting rhizobacteria. Encyclopedia of Genetics. Brenner,
polypeptides contain conserved glutamate and tyrosine S. and J.F. Miller, (Eds in chief) Academic Press:
residues (positions 15 and 19, respectively, in K. 1477-1480.
pneumoniae and the equivalents in A. calcoaceticus and M. r-5. Antoun, H., C. J. Beauchamp, N. Goussard, R. Chabot
and R. Lalande, 1998. Potential of Rhizobium and
extorquens). Those residues have been suggested previously Bradyrhizobium species as growth promoting bacteria on
as precursors in PQQ biosynthesis. Replacement of Glu-16 non-legumes: effect on radishes (Raphanus sativus L.). Plant
Soil 204: 57–67
by Asp and Tyr-20 by Phe in A. calcoaceticus PqqIV r-6. Azcon-Aguilar, C., V. Gianinazzi-Pearson, J. F.
abolished PQQ biosynthesis. A frame shift in K. Fardeau, S. Gianinazzi, 1986. Effect of vesicular-arbuscular
pneumoniae pqqA had the same result (Melulenberg et al., mycorrhizal fungi and phosphate solubilizing bacteria on
growth and nutrition of soybean in a neutral calcareous soil
1992). It was suggested that the PqqA/PqqIV polypeptide amended with 32P-45Ca-tricalcium phosphate. Plant and
might act as a precursor in PQQ biosynthesis (Goosen 1989; Soil. 96: 3–15.
r-7.
Goosen 1992; Melulenberg et al., 1992). r-8. Alghazali, R., S. H. M. Kmuhammad and Al-gzawl,
1986. Some observations on P solubilization by aerobic
In what way could a 24-amino-acid polypeptide be involved microorganisms isolated d from sediments of Al Khair River
in PQQ synthesis? Its small size makes a direct enzymatic Baghdad (Iraq). J Bio. Sci.Res.47:157-72.
r-9. Anderson, G., 1980. Assessing organic phosphorus in
function in the conversion of glutamate and tyrosine to PQQ soils. In The Role of Phosphorus in Agriculture (Khasawneh,
unlikely. A regulatory role of the gene IV product in the F.E., E.C. Sample and E.J. Kamprath, eds). Madison, USA:
expression of the other pqq genes is also improbable, since Amer. Soc. Agron. 411-432.
r-10.
previous experiments already showed that gene IV is also r-11. Bartel, B. 1997. Auxin biosynthesis. Ann. Rev. Plant
essential for PQQ synthesis in an E. coli strain. In these Physiol. Plant Mol. Biol. 48:51–66.
r-12. Belimov, A. A., A. P., .Kojemiakov and C. V.,
experiments, the expression of the pqq genes from A. Chuvarliyeva, 1995. Interaction between barley and mixed
calcoaceticus was under control of the E. coli lac promoter, cultures of nitrogen fixing and phosphate-solubilizing
which excludes a transcriptional control of these genes by bacteria. Plant Soil. 173:29–37.
r-13. Babu-Khan, S., T.C Yeo, W.L. Martin, M.R. Duron,
the gene IV peptide. Therefore, it is likely that the small R.D. Rogers and A.H. Goldstein, 1995. Cloning of a mineral
polypeptide has a more direct role in synthesis of the phosphate-solubilizing gene from Pseudomonas cepacia.
Appl. Environ. Microbiol. 61: 972-978.
coenzyme (Velterop1995). r-14.
r-15. Banik, S. and B. K. Dey, 1982. Available phosphate
Seven genes, called pqq genes, are required for PQQ content of an alluvial soil is influenced by inoculation of
biosynthesis in M. extorquens AM1, but their functions are some isolated phosphate solubilizing microorganisms Plant
Soil. 69: 353-364.
unknown (Moris et al., 1994; Nunn and Lidstrom 1986a, r-16. Barber, S.A., 1984. Soil Nutrient Bioavailability: A
Nunn and Lidstrom 1986b). The pqqD gene encodes a small Mechanistic Approach. John Wiley, New York
r-17. Beever, R. E. and D. J. W., Burns, 1980. Phosphorus
polypeptide of 29 amino acids containing conserved tyrosine uptake, storage and utilization by fungi. Advances in
and glutamate residues separated by three amino acids Botanical Research. 8:127-219.
(Moris et al., 1994). Tyrosine and glutamate have been r-18. Bossier,P,PM, Hofte and Verstrate1988.Adv.Microbial
Eco.10:385-413.
shown to be the precursors of PQQ biosynthesis, and it has r-19. Burgstaller, W. and F. Schinner, 1993. Leaching of
been proposed that the peptide might serve as the substrate metals with fungi. J. Biotechnol. 27: 91-116.
r-20. Chabot, R., C. J. Beauchamp, J. W. Kloepper and H.
for PQQ biosynthesis (Goosen, 1992; Houck, 1991; Antoun, 1998. Effect of phosphorous on root colonization
Meulenberg1992). and growth promotion of maize by bioluminescent mutants
of phosphate solubilizing Rhizobium leguminosarum biovar.
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Author Information
Nuzhat Ahmed
Centre for Molecular Genetics, University of Karachi
Sadaf Shahab
Centre for Molecular Genetics, University of Karachi
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