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Effects of protein-calorie malnutrition during suckling and post-weaning periods on discontinuous cranial traits in rats.

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AMERICAN JOURNAL OF PHYSICAL AN"HR0POLOGY 60:426430(1983)
Effects of Protein-Calorie Malnutrition During Suckling and
Post-Weaning Periods on Discontinuous Cranial Traits in Rats
SILVIA L. DAHINTEN AND HlbXOR M. PUCCIARELLI
Laboratorio de Investigacwnes Morfol6gicas, C&te&a IIIa de AnatomfG
Facultad de Ciencias Medicas, Universidad Nacwnul de La Plata, B u m s
Aires,Argentina
KEY WORDS
Discontinuous traits, Protein-calorie malnutrition,
Rat skull
ABSTRACT The influences of proteincalorie malnutrition 0,
and sex
during lactation and post-lactation on the frequencies of 25 discontinuous
cranial traits 0,
were investigated in Holtzman rats. Significant differences were observed in about 20% of the traits. Those traits were: the interfrontal fusion, the posterior incurvation of the palatine border, the double
maxillary foramen, the double posterior palatine foramen, and the double
frontal foramen. Total PCM was the nutritional fador which showed the
greatest influence on the variability of the DCTs. It was followed, in decreasing
order, by the PCM imposed during post-lactation and lactation. Sex had more
influence than early PCM but less than late PCM.It is concluded that despite
their apparent stability, a substantial number of DCTs were altered by both
biological (like sex) and environmental factors (like nutritional deficiencies)
imposed at different stages of postnatal development.
Discontinuous cranial traits (DCTs) are
bone formations that may appear in man
(Ossenberg, 1970; Corruccini, 1974a; Berry,
1975; Finnegan, 1978; Cheverud, Buikstra
and Twichell, 1979; Perizonius, 1979a,b),
nonhuman mammals Oeol, 1955; Deol and
Truslove, 1957; Berry and Searle, 1963;
Wickramaratne, 1974; Berry, 1977, 1978;
Cheverud and Buikstra, 1978),and other vertebrates (Grewal, 1978). Those traits are frequently used for human taxonomical studies
(Pucciarelli, 1971; Corruccini, 1972, 197413;
Berry and Berry, 1972; Sjdvold, 1973; Berry,
1974;Finnegan and Cooprider, 1978)because
their phenotypic expression is fairly independent of environmental stimuli and, in general, they possess Mendelian heritability.
Many studies showed that a nutritional deficiency brings about ossification disturbances in different mammals mimes, 1978).
Such disturbances would also alter the normal frequency of discontinuous skeletal
traits. A recent study mahinten and Pucciarelli, 1981)revealed that a nutritional deficiency imposed on weanling rats modified
the normal phenotypic expression of some
DCTs.
0 1983 ALAN R.LISS, INC.
The objective of the present work was to
determine whether a more acute and/or prolonged nutritional deficiency, such as a proteincalorie malnutrition during lactation and
post-lactation, could increase the nongenetic
variation of some DCTs in the growing rat.
The individual response of each sex was also
evaluated.
MATEXIAL AND METHODS
Pregnant Holtzman rats were kept in individual cages and fed water and stock diet ad
libitum until delivering. Thereafter, they
were subjected to either a control (C) or a
restricted (R) treatment. Control mothers
were fed stock diet ad libitum until weaning.
Their food intake was recorded daily. Restricted mothers were given only 50% of the
stock diet eaten daily by their respective controls.
At delivery, each litter was reduced to four
males and four females. At weaniq-21 days
of age-the offspring were allocated to one of
the followingtreatments:
Received October 6,1981, accepted September 27,1982.
426
S.L. DAHINTEN AND H.M.PUCCIARELLI
Control (CC): pups from control mothers
were fed a semisynthetic diet containing 28%
protein ad libitum.
Early malnutrition (RC): pups from restricted mothers were fed the same diet as
the CC rats.
Late malnutrition (CM): pups from control
mothers were fed a low-protein semisynthetic diet (6%protein) ad libitum.
pups from reTotal malnutrition 0:
stricted mothers were given the same diet as
CM.
The semisynthetic diets were prepared
weekly and according to the procedures developed by Dahinten and Pucciarelli (1981).
Diets were dried to constant weight and refrigerated until use. In this way, their quality remained unaltered and similar consistencies for both control and low-protein diets
were obtained. Body weights of both mothers, from delivery to weaning, and offspring,
until they were 49 days of age, were recorded
weekly. When the latter were 49 days old
they were killed with ether; their skulls were
cleaned following the technique of Luther
(Searle, 1954a)and stained with alizarin-red
“S”(Noback and Noback, 1944).
Twenty five DCTd were observed on 128
skulls (Table 1) under a dissecting microscope (20x1. Bilateral traits were observed
on both sides of the skull. All determinations
were made by the same observer, who was
not aware of the experimental group each
skull belonged to. Another observer verified,
on randomly chosen skulls, the determinations of the former. The sex and nutritional
factors (Tables 2-4) were obtained from comparisons between treatments. The statistical
analysis of the data consisted of z-tests between proportions; it was performed at the
Centro de Estudios Superiores para el Procesamiento de la Lnformacih (CESPI).
RESULTS
Weight differencesbetween the control and
restricted groups were observed from the
fourteenth day onwards. The recuperation
TABLE 1. Composition of the sample
Treatment
Control (CC)
Earlymalnutrition(RC)
LatemalnutritionCM)
Totalmalnutrition(RM)
Total
Males
Females
Symbol n Symbol n Total
MCC
MRC
MCM
MRM
FCC
12 FRC
18 FCM
18 FRM
62
14
15 29
10 22
22 40
19 37
66 128
TABLE 2. Comparison of the incidence of crania with
traits affected by sex between different groups of control
and stressed rats
Comparison
Proportion
z Test
Posterior incurvation of the left palatine border
-2.08*
0.58 - 0.93
MCC - FCC
-2.24*
0.60 - 1.00
MRC - FRC
-2.84**
0.62 - 0.96
MCM - FCM
2.41*
MRM-FRM
0.94 - 0.60
Left double frontal foramen
0.58 - 1.00
2.69**
MCC - FCC
0.73 - 0.70
n.8.
MRC - FRC
0.67 - 0.75
n.8.
MCM - FCM
MRM - FRM
0.78 - 0.75
n.s.
Right double frontal foramen
0.64 - 0.67
n.s.
MCC - FCC
MRC- FRC
0.90 - 0.80
n.8.
MCM - FCM
0.67 - 0.83
-2.24*
0.47 - 0.47
n.8.
MRM - FRM
Left double posterior palatine foramen
0.33 - 0.78
-2.33*
MCC - FCC
0.70 - 0.70
n.8.
MRC
.
._- FRC
MCM- FCM
0.71 - 0.67
n.8.
n.8.
MRM-FRM
0.71 - 0.50
~
~~
*P < 0.05; +*P< 0.01.
after weaning was not strong enough to raise
body weights to values similar to those of the
controls. The late and total-malnourished
groups did not increase in weight (Fig. 1).
Sex affected the posterior incurvation of
the left palatine border, and the double frontal and the left double posterior palatine foramina (Table 2). Nutrition affected the
posterior incurvation of the palatine border,
the interfrontal fusion, the double frontal foramen, the double maxillary foramen, and the
double posterior palatine foramen (Table 3).
Interactions between sex and nutrition were
found for the posterior incurvation of the palatine border, the interfrontal fusion, the double frontal foramen, the double maxillary
foramen and the right double posterior palatine foramen (Table 4).
‘The following discontinuous cranial traits were observed:
double frontal foramen (Berry and Searle, 1963); double maxillary foramen (Berry and Searle, 1963);double posterior palatine
foramen (Berry and Searle, 1963);mid-sphenoidal foramen (Deol
and Truslove, 1957); double oval foramen (Deol, 1955); double
hypoglossi foramen (Deol, 1955); double mental foramen -1,
1955); bentnose (Berry and Searle, 1963); incurvation of the
palatine border (Deol, 1955); interfrontal bone (Berry and Searle,
1963); bregmatic wormian bone (Pucciarelli, 1974); lambdoid
wormian bone (Pueeiarelli, 1974); third molar missing (Berry
and Searle, 1963); parted frontals (Berry and Searle, 1963);bone
reaorption in the eminentia incisiva of the mandible (Deol, 1955);
interfrontal fusion (Deol and h l o v e , 1957); internasal fusion
(Berry and Searle, 1963); squamosal-frontal fusion (Berry and
Searle. 1963);squamosal-parietalfusion (Berry and Searle, 1963);
occipital-periotic fusion (Berry and Searle, 1963); interparim
occipital fusion (Deol and Truslove, 1957); basi-preesphenoid fusion (Berry and Searle, 1963);basioccipital-basiesphenoidfusion
(Berry and Searle, 1963); basioccipital foramen present and
sphenoidal fissure.
EFFECTS OF F'CM ON DISCONTINUOUS TRAITS
Ol
7
12
27
28
35
45
49
AGE [DAYS I
Fig. 1. Increments in body weight in controls (uppertipped triangles); early malnourished (circles); late malnourished(squares); and total malnourished(down-tipped
triangles)groups.White: males; black females.
The traits which remained unaltered were:
the double hypoglossi foramen, the double
oval foramen, the bent-nose, the mid-sphenoidal foramen, the bregmatic wormian bone,
the parted frontals, and the sphenoid-occipital fissure. The following traits were absent:
the third molar missing, the squamosal-parietal fusion, the squamosal-frontalfusion, the
occipital-periotic fusion, the double mental
foramen, the bone resorption in the eminentia incisiva of the mandible, the internasal
fusion, the interparieto-occipital fusion, the
basi-preesphenoid fusion, the basiesphenoidbasioccipital fusion, the interfrontal bone,
and the basioccipital foramen.
DISCUSSION
About 20% of the measured trails were influenced by either PCM or PCM and sex.
PCM modified all of those traits whereas sex
affected only 2% of them. Although early
PCM had less influence than sex, its effect
was increased when it acted together with
late PCM (total PCM). In fact, the relative
427
effects of both sex and nutrition were: total
PCM > late PCM > sex > early PCM.
The major implications of those effects
were: a) not all of the DCTs are insensitive
to the influence of biological and/or environmental factors. On the contrary, there are
discontinuoustraits for which their variation
depends upon one or more of these fadors,
acting either per se or interactively; b) traits
apparently as stable as the DCTs may be
altered by a nutritional stress to an extent
greater than that evoked by genetically controlled fadors; and c) the DCTs seem to be
more sensitive to protracted nutritional deficits Qateand total PCM)than to acute deficiencies, such as those imposed during the
early period of postnatal development (early
PO.
The DCT which showed the greatest variation was the posterior incurvation of the left
palatine border. It was followed, in decreasing order of variation, by the interfrontal
fusion, the double frontal foramen, the double maxillary foramen, the posterior incurvation of the right palatine border, and the
double posterior palatine foramen. In addition, some of the traits, e.g., the double maxillary foramen and the interfrontal fusion,
showed specific nutritional variabilities as
Pucciarelli (1980)found for traits having continuous variation. The double frontal foramen showed predominant sexual variability.
The behavior of the other affected traits, i.e.,
the posterior incurvation of the palatine border and the double posterior palatine foramen, was not coherent because the variation
of one side was independent from that of the
opposite one.
The concept of specific nutritional variability is in agreement with Searle (1954b) and
Deol and Truslove (19571, who found nutritionally evoked changes in the discontinuous
traits of the mouse. The behavior of the inter.
frontal fusion concurs with both the decrease
of its frequency under conditions of a PCM
imposed during post-lactancy (Dahinten and
Pucciarelli, 1981)and the lower frequency of
the interparieto-occipital fusion, as observed
in mice born from mothers malnourished
during pregnancy and lactation (Deol and
Truslove, 1957).The higher frequenciesfound
by Deol and Truslove in oval and mid-sphenoidal foramina and the lower frequencies in
the posterior incurvation of the palatine border are at variance with those observed in
this experiment, presumably because of the
different kind of diets and animals employed.
S.L. DAHINTEN AND H.M. PUCCIARELLI
428
TABLE 3. Comparison of the incidence of crania with
traits affected by nutrition between different groups of
control and stressed rats
Comparison
Proportion
Factor
z Test
TABLE 3. Continued
Comparison
Proportion
Factor
z Test
Posterior incurvation of the left palatine border
MCC - MRC
0.58 - 0.60 Early PCM
n.s.
0.62 - 0.94
Early PCM -2.32*
MCM - MRM
2.93*
FCM - FRM
0.96 - 0.60 Early PCM
n.8.
0.93 - 1.00 Early PCM
FCC - FRC
n.s.
0.58 - 0.62
Late PCM
MCC - MCM
n.s.
0.93 - 0.96
Late PCM
FCC - FCM
0.60 - 0.94
Late PCM -2.20*
MRC - MRM
Late PCM
2.34*
FRC - FRM
1.00 - 0.60
0.58 - 0.94
Total PCM -2.34*
MCC - MRM
2.14*
0.93 - 0.60
Total PCM
FCC .FRM
Posterior incurvation of the right palatine border
n.s.
0.92 - 0.70 Early PCM
MCC - MRC
n.6.
0.93 - 0.90 Early PCM
FCC FRC
n.s.
0.95 - 1.00 Early PCM
MCM - MRM
Early PCM
n.s.
FCM - FRM
0.96 - 0.95
Late PCM
n.8.
MCC -MCM
0.92 - 0.95
n.s.
0.93 - 0.96
Late PCM
FCC - FCM
0.70 - 1.00
Late PCM -2.40*
MRC - MRM
Late PCM
n.8.
0.90 - 0.95
FRC - FRM
n.8.
0.92 - 1.00
Total PCM
MCC - MRM
Total PCM
n.s.
FCC - FRM
0.93 - 0.95
Interfrontal fusion
0.25 - 0.02 Early PCM
n.8.
MCC - MRC
0.21 - 0.10 Early PCM
n.8.
FCC - FRC
0.05 - 0.00 Earl; PCM
n.s.
MCM - MRM
n.8.
FCM - FRM
0.00 - 0.00 Early PCM
n.s.
Late PCM
0.25 - 0.05
MCC MCM
2.36*
Late PCM
FCC FCM
0.21 - 0.00
n.8.
Late PCM
0.02 - 0.00
MRC - MRM
n.8.
0.10 - 0.00
Late PCM
FRC - FRM
2.18*
0.25 - 0.00
Total PCM
MCC - MRM
2.17*
Total PCM
FCC - FRM
0.21 - 0.00
Left double frontal foramen
MCC
MRC
0.58 - 0.73
Earlv PCM
n.s.
-- - .
-. FCC - FRC
i.00 - 0.70 Earl; PCM 2.19*
n.6.
0.67 - 0.78
Early PCM
MCM - MRM
Early PCM
n.s.
FCM - FRM
0.75 - 0.75
n.s.
0.58 - 0.67
Late PCM
MCC - MCM
2.04*
1.00 - 0.75
Late PCM
FCC - FCM
n.8.
0.73 - 0.78
Late PCM
MRC - MRM
Late PCM
n.6.
0.70 - 0.75
FRC - FRM
n.8.
Total PCM
0.58 - 0.78
MCC - MRM
2.03*
TotalPCM
1.00 - 0.75
FCC - FRM
Right double frontal foramen
n.s.
0.64 - 0.90 Early PCM
MCC - MRC
0.67 - 0.80 Early PCM
n.s.
FCC - FRC
Early PCM
n.8.
0.67 - 0.47
MCM MRM
0.83 - 0.47
FCM - FRM
n.s.
Early PCM
0.64 - 0.67
n.8.
Late PCM
MCC .MCM
0.67 - 0.83
Late PCM
n.8.
FCC - FCM
0.90 - 0.47
Late PCM
2.23 *
MRC - MRM
n.6.
0.80 - 0.47
Late PCM
FRC - FRM
0.64 - 0.47
Total PCM
n.8.
MCC - MRM
Total PCM
n.8.
FCC - FRM
0.67 - 0.47
Left double maxillary foramen
0.33 - 0.40 Early PCM
n.8.
MCC - MRC
n.8.
FCC - FRC
0.36 - 0.70 Early PCM
n.8.
MCM - MRM
0.24 - 0.41 Early PCM
0.08 - 0.40 Early PCM -2.50*
FCM - FRM
n.8.
0.33 - 0.24
MCC - MCM
Late PCM
0.36 - 0.08
Late PCM
2.10*
FCC - FCM
0.40 - 0.41
Late PCM
n.s.
MRC - MRM
Late PCM
n.8.
FRC - FRM
0.70 - 0.40
Total PCM
n.8.
0.33 - 0.41
MCC - MRM
Total PCM
n.s.
0.36 - 0.40
FCC - FRM
Right double maxillary foramen
n.8.
0.33 - 0.10 Early PCM
MCC- MRC
2.39*
0.43 - 0.00 Early PCM
FCC FRC
n.8.
MCM - MRM
0.24 - 0.35 Early PCM
Earh PCM
n.8.
FCM - FRM
0.37 - 0.47
Late PCM
n.8.
0.33 - 0.24
MCC - MCM
Late PCM
n.8.
0.43 - 0.37
FCC - FCM
Late PCM
n.8.
0.10 - 0.35
MRC - MRM
Late PCM -2.92*
FRC - FRM
0.00 - 0.47
n.6.
0.33 - 0.35
Total PCM
MCC - MRM
n.8.
FCC .FRM
0.43 - 0.47
Total PCM
Left double posterior palatine foramen
n.8.
0.33 - 0.70 Early PCM
MCC- MRC
n.8.
FCC - FRC
0.78 - 0.70 Early PCM
n.8.
0.71 - 0.70 Early PCM
MCM - MRM
n.s.
0.67 - 0.50 Early PCM
FCM - FRM
Late PCM -2.13*
0.33 - 0.71
MCC - MCM
Late PCM
n.s.
0.78 - 0.67
FCC - FCM
n.8.
0.70 - 0.70
Late PCM
MRC - MRM
n.8.
0.70 - 0.50
Late PCM
FRC - FRM
0.33 - 0.70
Total PCM -2.00*
MCC - MRM
n.8.
Total PCM
FCC - FRM
0.78 - 0.50
Right double posterior palatine foramen
n.8.
0.75 - 0.70 Early PCM
MCC - MRC
n.8.
FCC - FRC
0.50 - 0.60 Early PCM
Earl; PCM
n.8.
MCM-MRM
0.57-0.29
n.8.
0.62 - 0.60 Early PCM
FCM - FRM
Late PCM
n.8.
MCC - MCM
0.75 - 0.57
Late PCM
n.8.
FCC - FCM
0.50 - 0.62
2.05*
0.70 - 0.29
Late PCM
MRC - MRM
n.8.
0.60 - 0.60
Late PCM
FRC - FRM
MCC - MRM
0.75 - 0.29
Total PCM
2.42*
FCC - FRM
0.50 - 0.60
Total PCM
n.8.
*P 6 0.05; **P4 0.01.
Berry and Berry (1972) stated that discontinuous traits are independent from biological fadors. However, the present observations in the rat along with those of Howe and
Parsons (1967) in the mouse and Conuccini
(1974a,b) in man suggest the existence of
strong influences of biological factors, such
as age and sex, on discontinuous traits. Indeed, the interaction between sex and PCM
was quite evident since in eight out of nine
of the affected traits there was a comparison
for which the combination of both fadors
evoked significant differences. This interaction was made evident by the behavior of the
left posterior palatine border, which suggests
that a nutritional stress may affect males
and females differently.
While the influence of sex may be subjected
to epigenetic polymorphism (Berry and
Searle, 19631,the nutritional influences must
~
~
~
~
~
429
EFFECTS OF PCM ON DISCONTINUOUS !I'RAl"s
TABLE 4. Comparison of the incidence of crania with
traits affected by sex-nutrition interaction between
differentgroups of control and stressed rats
Comparison
Proportion
Factors
z Test
TABLE 4. Continued
Comparison
Proportion
Factors
z Test
Posterior incurvation of the left palatine border
MCC - FRC 0.58 - 1.00 S + early PCM -2.32*
FCC - MRC 0.93 - 0.60 S + early PCM
n.6.
MCM - FRM 0.62 - 0.60 S + early PCM
n.8.
FCM - MRM 0.96 - 0.94 S + early PCM
n.8.
MCC - FCM 0.58 - 0.96 S + late PCM -2.85*
FCC - MCM 0.93 - 0.62 S + late PCM
2.05*
MRC - FRM 0.60 - 0.60 S + late PCM
n.8.
FRC - MRM 1.00 - 0.94 S + late PCM
n.8.
MCC - FRM 0.58 - 0.60 S + total PCM
n.8.
FCC - MRM 0.93 - 0.94 S + total PCM
n.8.
Posterior incurvation of the right palatine border
MCC - FRC 0.92 - 0.90 S + early PCM
n.8.
FCC - MRC 0.93 - 0.70 S + early PCM
n.8.
MCM - FRM 0.95 - 0.95 S + earlv PCM
n.8.
FCM - MRM 0.96 - 1.00 s + early PCM
n.8.
MCC - FCM 0.92 - 0.96 S + late PCM
n.8.
FCC - MCM 0.93 - 0.95 S + late PCM
n.8.
MRC - FRM 0.70 - 0.95 S + late PCM -1.90*
FRC - MRM 0.90 - 1.00 S + late PCM
n.8.
MCC - FRM 0.92 - 0.95 S + total PCM
n.8.
FCC - MRM 0.93 - 1.00 S + total PCM
n.8.
Interfrontal fusion
MCC - FRC 0.25 - 0.10 S + early PCM
n.8.
FCC MRC
0.21 - 0.02 S + early PCM
n.8.
MCM - FRM 0.05 - 0.00 S + early PCM
n.8.
FCM - MRM 0.00 - 0.00 S + early PCM
n.8.
MCC - FCM 0.25 - 0.00 S + late PCM
2.56*
FCC - MCM 0.21 - 0.05 S + late PCM
n.8.
MRC - FRM 0.02 - 0.00 S + late PCM
n.8.
FRC - MRM 0.10 - 0.00 S + late PCM
n.8.
MCC - FRM 0.25 - 0.00 S + total PCM
2.35*
FCC - MRM 0.21 - 0.00 S + total PCM
2.01*
Left double frontal foramen
MCC - FRC 0.58 - 0.70 S + early PCM
n.8.
FCC - MRC
1.00 - 0.73 S + early PCM
n.8.
MCM - FRM 0.67 - 0.75 S + early PCM
n.8.
FCM - MRM 0.75 - 0.78 S + early PCM
n.8.
MCC - FCM 0.58 - 0.75 S + late PCM
n.8.
FCC - MCM 1.00 - 0.67 S + late PCM
2.42*
MRC - FRM 0.73 - 0.75 S + late PCM
n.8.
FRC - MRM 0.70 - 0.78 S + late PCM
n.8.
MCC - FRM 0.58 - 0.75 S + total PCM
n.8.
FCC - MRM 1.00 - 0.78 S + total PCM
n.8.
Right double frontal foramen
n.8.
0.64 - 0.80 S + early PCM
MCC - FRC
n.8.
FCC - MRC 0.67 - 0.90 S + early PCM
n.8.
MCM - FRM 0.67 - 0.47 S + early PCM
2.67*
FCM - MRM 0.83 - 0.47 S + early PCM
MCC- FCM
0.64 - 0.83 S + late PCM
n.8.
n.8.
FCC - MCM 0.67 - 0.67 S + late PCM
2.37*
MRC - FRM 0.90 - 0.47 S + late PCM
n.8.
FRC - MRM 0.80 - 0.47 S + late PCM
n.8.
MCC - FRM 0.64 - 0.47 S + total PCM
n.8.
FCC-MRM 0.67 - 0.47 S + total PCM
Left double maxillary foramc!n
n.8.
MCC - FRC 0.33 - 0.70 S + early PCM
n.8.
FCC - MRC 0.36 - 0.40 S + early PCM
n.8.
MCM - FRM 0.24 - 0.40 S + early PCM
S
+
early
PCM
-2.50*
FCM - MRM 0.0s - 0.41
S
+
late
PCM
n.8.
MCC - FCM 0.33 - 0.08
FCC
0.24 S + late PCM
n.8.
~- - MCM
. _ .0.36
~- ~
n.8.
MRC - FRM 0.40 - 0.40 S + late PCM
n.8.
FRC - MRM 0.70 - 0.41 S + late PCM
n.8.
MCC - FRM 0.33 - 0.40 S + total PCM
n.8.
FCC - MRM 0.36 - 0.41 S + total PCM
Right double maxillary foramen
2.01*
MCC - FRC 0.33 - 0.00 S + early PCM
n.s.
FCC - MRC 0.43 - 0.10 S + early PCM
n.8.
MCM - FRM 0.24 - 0.47 S + early PCM
n.8.
FCM - MRM 0.37 - 0.35 S + early PCM
n.8.
FCC - MCM 0.43 - 0.24 S + late PCM
n.8.
MCC - FCM 0.33 - 0.37 S + late PCM
n.s.
MRC - FRM 0.10 - 0.47 S + late PCM
-2.13*
FRC - MRM 0.00 - 0.35 S + late PCM
n.8.
MCC - FRM 0.33 - 0.47 S + total PCM
n.8.
FCC MRM 0.43 - 0.35 S + total PCM
Right double posterior palatine foramen
n.8.
MCC - FRC 0.75 - 0.60 S + early PCM
n.6.
FCC - MRC 0.50 - 0.70 S + early PCM
n.8.
MCM- FRM 0.57 - 0.60 S + early PCM
2.09*
FCM MRM 0.62 - 0.29 S + early PCM
n.8.
MCC - FCM 0.75 - 0.62 S + late PCM
n.8.
FCC - MCM 0.50 - 0.57 S + late PCM
n.s.
MRC - FRM 0.70 - 0.60 S + late PCM
n.8.
FRC .MRM 0.60 - 0.29 S + late PCM
n.8.
MCC - FRM 0.75 - 0.60 S + total PCM
n.8.
FCC - MRM 0.50 - 0.29 S + total PCM
be explained by other factors. The behavior
of the interfrontal fusion under PCM can be
readily explained by other factors. The behavior of the interfrontal fusion under PCM
can be readily explained because it is a hyperostotic trait, i.e., its ossification increases
with age (Ossenberg, 1970). The delay in ossification brought about by PCM partly arrested the process of cranial sinostosis, and
hence led to the persistence of the interfrontal suture. Because the foraminae are not
hyperostotic traits, their variation might be
due to their functional relation with the
nerves and vessels which pass them through.
The degree of branching of the blood vessels
and/or nerve fibers which pass through a foramen determine its simple or multiple character. PCM might have modified that degree
of branching, and given rise to the observed
foraminal variation.
~
~
-
~
*P c 0.05; **P< 0.01.
S :sex.
CONCLUSIONS
DCTs do not show an even behavior. Although most are stable a substantial number
430
S.L. DAHINTEN AND H.M. PUCCJARELLI
are not, 80 that they are prone to change with
biological factors, such as sex, andlor environmental ones, such as nutrition.
It is possible that there is not always reciprocity between a discontinuous trait in the
rat skull and a similar trait in the human
skull but the behavior shown for the DCTs of
the rat represents a biological basis from
which the presumptive behavior of human
DCTs can be inferred. Within that conceptual framework, however, the results of the
present study are relevant to physical anthropology.
Despite the taxonomic distance between
humans and rodents, anthropologists must
be aware that the differential behavior of the
DCTs cautions against the careless use of
those traits as parameters for racial classification. However, the use of traits with a specific nutritional component can be fruitful for
other studies, e.g., the osseous modifications
in populations subjected to a nutritionally
deficient environment.
ACKNOWLEDGMENTS
The authors are grateful to Dr. Mario H.
Niveiro (C6tedra IIIa de Anatomia; Facultad
de Ciencias M6dicas; Universidad Nacional
de La Plata), Ing. H&r D. Ginzo (Centro de
Ecofisiologia Vegetal), and Lic. Evelia E.
Oyhenart, and Mrs. Maria C. Muiie (Laboratori0 de Investigaciones Morfol6gicas) for
their highly valuable assistance; and to Refinerias de Maiz S.A.I.C.E., Celulosa Argentina S.A., and Laboratorios Ruminal S.A.I.C.
for the economic support.
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