Advances in Bioscience and Biotechnology, 2011, 2, 27-32 ABB
doi:10.4236/abb.2011.21005 Published Online February 2011 (http://www.SciRP.org/journal/abb/).
Published Online February 2011 in SciRes. http://www.scirp.org/journal/ABB
Genetic variability of the Saudi Arabian Uromastyx aegyptia
microlepis using protein and isoenzymes electrophoreses
Sayed Amin Mohamed Amer1,2
1Biotechnology Department, Faculty of Science, Taif University, Saudi Arabia;
2Zoology Department, Faculty of Science, Cairo University, Egypt.
Received 24 December 2010; revised 29 January 2011; accepted 15 February 2011.
ABSTRACT
Electrophoresis for SDS-proteins and isoenzymes
were conducted to investigate the genetic variations
within the agamid lizard Uromastyx aegyptia microle-
pis inhabiting the desert of Saudi Arabia. Samples
were collected from four localities: a) Ushayrah near
the town of Taif, b) Al Gwaih near Riyadh, c) Nai-
ryah near Dammam and d) Mouileh near Tabok. A
range of 7 to 14 protein bands were recorded in the
patterns of the studied samples as measured by
SDS-polyacrylamide gel electrophoresis. Among them,
only one fraction was recorded in all samples as a
common protein band. Six arbitrary chosen enzymes
were examined by native-polyacrylamide gel elec-
trophoresis. They were α and β esterase (Est), acid
phosphatase (Acph), Alcohol dehydrogenase (Adh),
Aldehyde oxidase (Ao) and peroxidase (Px). Seven-
teen heterogeneous alleles have been recorded; seven
of them were fixed in all populations and 10 were
polymorphic. Nearly all recorded alleles were mono-
meric in all samples. α-Est2, β-Est2, Acph2 and Px1
were restricted to Tabok samples and were not re-
corded in other localities. β-Est3, Acph3, Adh1,
Adh1and Px2 were not recorded in Taif samples and
the latter one was not recorded in the Dammam sam-
ples. The similarity coefficient that has been calcu-
lated according to the number of sharing bands indi-
cated the clustering of Tabok and Dammam popula-
tions together. The constructed tree based on the
sharing protein bands and isozyme alleles revealed
similar results regarding the kinship of both popula-
tions. The present results indicated that the popula-
tions of this subspecies exhibits high genetic variabil-
ity among its populati ons .
Keywords: Uromastyx; Isoenzymes; Population
Genetics; Arabia
1. INTRODUCTION
Spiny-tailed Uromastyx are small to medium-sized,
ground-or rock-dwelling agamid lizards. The animals
have a bulky, depressed body and strong, short limbs.
The tail is covered by spiny scales, arranged in distinct
whorls. Uromastyx habitats are generally characterized
by high temperatures, low precipitation, sparse vegeta-
tion and marked seasonal changes. A detailed discussion
of the taxonomic history of the Uromastyx aegyptia
group is provided that is now recognized as three valid
species (U. aegyptia, U. leptieni and U. occidentalis) [1].
However, Uromastyx aegyptia has been recognized as a
polytypic species with three subspecies (aegyptia, lep-
tieni and microlepis) [2]. The main characters to distin-
guish the members of the Uromastyx aegyptia group
from other Uromastyx species groups are their large
body size combined with very small body scales. The
diagnostic features to distinguish the members of the
Uromastyx aegyptia group are the lack of enlarged tu-
bercular scales on the flanks and the presence of skin
folds at the sides of the neck covered with tubercles in U.
a. microlepis [3]. U. a. microlepis was also suggested to
be a valid subspecies by many authors [3-7]. It is found
in all Arabia and Iraq, commonly, amongst populations
of U.a. aegyptia. Their potential contact zone is at the
east of Wadi Araba (Jordan and Palestine) and at the east
of Wadi Sawawin in the Jebel as-Sinfa region (Tabok)
where members of both subspecies are found sympatri-
cally, while taxa from Eastern Arabia are exclusively
belonging to U. a. microlepis [3]. Recent molecular
studies investigated the systematics of Uromastyx [8-12]
but few biochemical studies have been conducted [12,5].
The present investigation aims to study the electropho-
retic behavior of several isoenzymes and proteins for U.
a. microlepis from different localities of its range in or-
der to address the genetic variability within this subspe-
cies.
2. MATERIALS AND METHODS
2.1. Samples
At least, three samples of U. a. microlepis were collected

S. A. M. Amer / Advances in Bioscience and Biotechnology 2 (2011) 27-32 28
from each of four localities encompassing the different
ecological habitats of the Saudi Arabian desert (Figure
1). These localities were Ushayrah, 70 km northeast to
Taif, AlGwaih 200 km west to Riyadh, Nairyah 200 km
west to Dammam and Mouileh 250 km Southwest to
Tabok.
Animals were taken to the lab, killed and dissected.
Samples of blood and muscle tissues were taken and
immediately frozen at - 80ºC.
2.2. SDS-Protein
SDS-polyacrylamide gel electrophoresis was performed
in 14 % acrylamide slab gels following the system of
[13]. Protein extraction was conducted by homogenizing
1.0 g of tissue in 2 ml saline solution NaCl (0.9%) using
a manual homogenizer. Samples were centrifuged at
5000 rpm for 10 min. The supernatants that contain pro-
tein extract were kept deep-frozen until use for electro-
phoresis analysis. A volume of 20 L protein extract was
added to 10 L of treatment buffer. 20 L of the mixture
was loaded in the gel. After electrophoresis, the gel was
stained by commassie brilliant blue. The gel was de-
stained after the appearance of bands and photographed.
2.3. Isoenzymes
Six isoenzymes were used in this study: α and β-esterase
(Est), acid phosphatase (Acph), alcohol dehydrogenase
(Adh), aldehyde oxidase (Ao) and peroxidase (Px).
Isoenzymes were electrophoresed in 10% native-poly-
acrylamide gel as described by Stegemann et al. [14].
For isoenzyme extraction, 1.0 g of tissue was homoge-
nized in 2 mL saline solution NaCl (0.9%) using a man-
ual homogenizer. The homogenates were centrifuged
Figure 1. A map with sample sites indicated (stars) and the
entire distribution (circled area) of Uromastyx aegyptia micro-
lepis.
at 5000 rpm for 10 minutes and the supernatants were
kept at -20ºC until use. For electrophoresis, 30 μL of the
extract was mixed with 10 L of treatment bufferand 35
μL of this mixture was applied to the well. In gel stain-
ing, protocols published for  and  -Est, Ao and Acph,
Adh and Px were used [15-18]. Gels were washed two or
three times with tap water; fixed in ethanol/ 20% glacial
acetic acid (9:11 v/v) for 24 hours and photographed.
2.4. Statistics
All gels of protein and isozyme electrophoresis were
scanned using Gel Doc-2001 Bio-Rad system. For
isoenzymes, the bands of enzyme activity were desig-
nated using the known system of nomenclature [19]. An
abbreviation which corresponds to the name of the en-
zyme designated each locus. When multiple loci were
involved, the fastest anodal protein band was designated
as locus one, the next as locus two and so on. A similar-
ity matrix of allele presence (1) and absence (0) was
constructed and the dendrogram was obtained by the
UPGMA method from NTSYS-pc package [20]. The
similarity coefficient was calculated [21,22] as follows:
2 (number of sharing bands)/ number of bands in popu-
lation A + number of bands in population B.
3. RESULTS AND DISCUSSION
The number of protein bands that were obtained from
this study varied between and within populations. A
range of 7 to 14 bands were recognized in the pattern of
the studied populations with minimum numbers in the
samples of Taif and Riyadh and maximum numbers in
Tabok (12 bands) and Dammam populations (12 bands)
as measured by SDS-polyacrylamide electrophoresis
(Table 1). The molecular weights of these bands showed
a wide range between a maximum of 89 kDa and a
minimum of 14 kDa. All samples studied recorded one
shared band with a molecular weight of 33.534 kDa.
There were no shared bands between populations at the
extremes of the protein profile. The number of shared
bands were 4, 9, 5 and 11 within Taif, Dammam, Riyadh
and Tabok populations, respectively. When the different
localities were compared, the number of shared bands
were 3 between Taif and Dammam populations, 2 be-
tween Taif and Riyadh populations, 4 between Taif and
Tabok populations, 4 between Dammam and Riyadh
populations, 9 between Dammam and Tabok populations
and 5 between Riyadh and Tabok populations. Three
unique bands were found in the Taif population at the
molecular weight of 89, 88 and 60 kDa and two unique
bands were shown in the Dammam population at 68 and
14 kDa. The electrophoretic banding of proteins indi-
cated that the populations of the present study have sim-
ilar bands within the molecular weight of the range be-

S. A. M. Amer / Advances in Bioscience and Biotechnology 2 (2011) 27-32 29
tween 73 and 63 kDa (as shown in Tab l e 1 ) and in the
range between 46 and 16 kDa. The former range is
comparable to vertebrate albumin (65 -70 kDa) [23,24]
that is considered as a good indicator for species differ-
entiation [25]. These similarities, therefore, could be
considered as an indication of gene flow among the stu-
died samples and do not refer to any fragmentation. The
similarity coefficients that were calculated according to
the number of sharing protein bands are shown in Table
2.
The Tabok population recorded the highest similarity
to the Dammam population with a coefficient of 0.69
and to the Riyadh population with a coefficient of 0.48.
The similarity coefficients between the Taif population
and each of the three other populations were 0.32, 0.29
and 0.38 for the Dammam, Riyadh and Tabok popula-
tions, respectively. The elctrophoretic behavior of the six
studied isoenzymes was shown as alleles and their rela-
tive mobility in Tables 3 to 8. Seventeen presumptive
heterogeneous alleles have been recorded in the present
study. Among the twelve microlepis samples studied
herein, the α and β-Est, Acph, Ao and Px isozymes re-
corded three putative genotypes and three allelic prod-
ucts for each (Tables 3-5, 7, 8). All genotypes were
identified as monomers (the complete enzyme consists
of only one polypeptide) with one allele. Ad h isoezyme
was identified by two genotypes with two monomeric
alleles (Table 6). Seven alleles for α-Est3, β-Est1, Acph1,
Adh2, Ao1, Ao3 and Px3 were shown to be fixed in all
samples. The samples from Tabok recorded the highest
diversity of phenotypes where α-Est2, β-Est2, Acph2 and
Px1 were characteristic to this population and were not
recorded in other populations. On the other hand, Taif
samples were shown to be the lowest in phenotypic di-
versity where β-Est3, Acph3, Adh1, Adh1 and Px2 were
not recorded in this population. The samples from the
vicinity of Tabok were collected from Mouileh (sandy
desert) which is located close to the Red Sea in the
northwest of the Kingdom and is found in sympatry with
U. a. aegyptia. On the other hands, Taif is located in the
Sarawat mountains (2000-3000 m altitude) with rocky
nature that could have exerted some effect on the genetic
variability of the population inhabiting this area.
The heterogeneity of the isozyme electrophoreses
among populations was also revealed in earlier investi-
gations. Esterase and alcohol dehydrogenase showed
four fractions in their loci and these fractions were pro-
posed to be obtained by the combinations of trimer po-
lypeptides [26]. A trimeric structure was reported for
esterase in human, pig, rat, and guinea pig [27]. Similar
to the present study, multiple forms were reported for
aldehyde oxidase in Uromastyx and other vertebrates
[12].
The constructed dendrogram (Figure 2(a)) showed
clustering of Tabok, Dammam and Riyadh populations.
The Taif population came basal in the dendrogram since
Table 2. The estimated similarity coefficients calculated ac-
cording to the shared protein bands among the studied popula-
tions.
TaifDammam Riyadh Tabok
Taif 0.32 0.29 0.38
Dammam 0.42 0.69
Riyadh 0.48
Table 1. Protein band molecular weights measured in kDa in samples of the different populations studied.
Taif (Ushayrah) Dammam (Nairyah) Riyadh (AlGwaih) Tabok (Mouileh)
MW Lane1 Lane2 Lane3 Lane4 Lane5 Lane6 Lane7 Lane8 Lane9
89.463 - - - - - - - - +
88.218 - - - - - - - - +
85.951 - - - + + - - + +
83.910 - - - + + - - + +
74.559 + + + + + - + + +
68.134 - - - - + - - - -
63.015 + + + - + + + + +
60.784 - - - - - - - - +
46.946 + + + + + + - + +
42.303 + - - + + - - + +
33.534 + + + + + + + + +
29.382 + - + + + + + + +
22.422 - + + + + + + + +
16.471 - + + + + + + + +
14.636 - - - + - - - + -

S. A. M. Amer / Advances in Bioscience and Biotechnology 2 (2011) 27-32 30
Ta b le 3. The recorded phenotypes and the relative mobility (RF) for the electrophoretic pattern of α-Est isoenzyme in the studied
populations of U. a. microlepis. Lanes are as follow: 1-4 (Taif samples), 5-6 (Dammam samples), 7-9 (Riyadh samples) and 10-12
(Tabok samples).
RF Alleles Lane1 Lane2 Lane3 Lane4 Lane5 Lane6Lane7Lane8Lane9Lane10 Lane11 Lane12
0.033 α-Est3 + + + + + + + + + + + +
0.890 α-Est2 - - - - - - - - - - + -
0.946 α-Est1 + + - - - - - - + + + +
Table 4. The recorded phenotypes and the relative mobility (RF) for the electrophoretic pattern of β-Est isoenzyme in the studied
populations of U. a. microlepis. Lanes are as follow: 1-4 (Taif samples), 5-6 (Dammam samples), 7-9 (Riyadh samples) and 10-12
(Tabok samples).
RF Alleles Lane1 Lane2 Lane3 Lane4 Lane5 Lane6Lane7Lane8Lane9 Lane10 Lane11 Lane12
0.024 β-Est3 - - - - + + - + - + + +
0.867 β-Est2 - - - - - - - - - - + -
0.917 β-Est1 + + + + + + + + + + + +
Ta ble 5. The recorded phenotypes and the relative mobility (RF) for the electrophoretic pattern of Acph isoenzyme in the studied
populations of U. a. microlepis. Lanes are as follow: 1-4 (Taif samples), 5-6 (Dammam samples), 7-9 (Riyadh samples) and 10-12
(Tabok samples).
RF Alleles Lane1 Lane2 Lane3 Lane4 Lane5 Lane6Lane7Lane8Lane9 Lane10 Lane11 Lane12
0.028 Acph3 - - - + + + - + - + + +
0.858 Acph1 - - - - - - - - - - + -
0.931 Acph1 + + + + + + + + + + + +
Table 6. The recorded phenotypes and the relative mobility (RF) for the electrophoretic pattern of Adh isoenzyme in the studied
populations of U. a. microlepis. Lanes are as follows: 1-4 (Taif samples), 5-6 (Dammam samples), 7-9 (Riyadh samples) and 10-12
(Tabok samples).
RF Alleles Lane1 Lane2 Lane3 Lane4 Lane5Lane6 Lane7Lane8 Lane9Lane10 Lane11 Lane12
0.029 Adh2 + + + + + + + + + + + +
0.061 Adh1 - - - + + + + + + + + +
Table 7. The recorded phenotypes and the relative mobility (RF) for the electrophoretic pattern of Ao isoenzyme in the studied popu-
lations of U. a. microlepis. Lanes are as follow: 1-4 (Taif samples), 5-6 (Dammam samples), 7-9 (Riyadh samples) and 10-12 (Tabok
samples).
RF Alleles Lane1 Lane2 Lane3 Lane4Lane5Lane6Lane7Lane8Lane9 Lane10 Lane11 Lane12
0.030 Ao 3 + + + + + + + + + + + +
0.060 Ao 2 - - - - + + + + + + + +
0.082 Ao 1 + + + + + + + + + + + +
Table 8. The recorded phenotypes and the relative mobility (RF) for the electrophoretic pattern of Px isoenzyme in the studied popu-
lations of U. a. microlepis. Lanes are as follow: 1-4 (Taif samples), 5-6 (Dammam samples), 7-9 (Riyadh samples) and 10-12 (Tabok
samples).
RF Alleles Lane1 Lane2 Lane3 Lane4 Lane5 Lane6Lane7Lane8Lane9Lane10 Lane11 Lane12
0.634 Px3 + + + + + + + + + + + +
0.830 Px2 - - - - - - + + + + + -
0.926 Px1 - - - - - - - - - - + -
it showed the lowest similarities to the other populations
(Ta bl e 2). The application of the UPGMA clustering on
the raw data obtained from isoenzymes and SDS pro-
teins split the populations (Figure 2(b)) into two main
clusters: the populations of Taif and AlGwaih (Riyadh)
as one group and the populations of Tabok and Dam-

S. A. M. Amer / Advances in Bioscience and Biotechnology 2 (2011) 27-32 31
(a)
(b)
Figure 2. Dendrogram based on similarity coefficients (A)
and UPGMA (B) analysis of genetic similarity obtained
from isozyme and protein data showing relationships
among the studied populations of U. a. microlepis.
mam as another group. By comparing this tree to that of
Figure 2(a), it was obvious that there was a concordance
regarding the kinship of Tabok and Dammam popula-
tions. The disconcordance between the two dendrograms
was found in the relationship of both Taif and Riyadh
populations. This could be attributed to the extremes in
the raw data applied for constructing Figure 2(b). Both
results indicated that U. a. microlepis have a wide ge-
netic variability between populations that are explained
by adaptations to different environmental conditions.
Uromastyx a. microlepis from Taif inhabits mountainous
areas and habitats with thick layers of stones and rocks
which are not suitable for this subspecies [28]. The
common habitats for other populations are the open ar-
eas with hard, diggable substrates like coarse sand, grav-
els and sparse vegetation [29].
4. CONCLUSION
I conclude that protein and isoenzyme electrophoreses is
a powerful tool in targeting the genetic variability within
the Uromastyx aegyptia microlepis populations. This
technique provides for further molecular investigations
that are necessary to address the genetic differentiation
of this subspecies more accurately.
5. ACKNOWLEDEMENTS
The Taif University is acknowledged for the financial support (grant
number 1-431-627). I thank the students of the biotechnology and
biology departments in the Faculty of Science for their assistance in
collecting animal samples from the wild. My thanks extend also to Dr.
Mahmoud Salman, a Canadian scientist for his assistance in reviewing
the manuscript grammatically.
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