An electrophoretic characterization of serum proteins of the collared peccary (Tayassu tajacu)

0305-0491/8zt $3.00 + 0.00 © 1984 Pergamon Press Ltd

, pp. 569 575, 1984

,OPHORETIC C H A R A C T )TEINS OF THE COLLA

ON OF ECARY

(TA YASSU TAJACU) ~ILLER, E. C. HELLGREN,L. W. VARN Department of Wildlife and Fisheries Sciences, Texas A&M University and *Agricultural Research and Extension Center, Texas A&M Uni

E. GRANT )n, TX 77843, USA ,', TX 78801, USA

(Receiced 14 March 1984) Akstraet--1. Serum proteins of the collared peccary ((Ta. Tayassu tajaci electrophoresis for wild adult males and females, nursing young, an( captivity. 2. Electrophoretic profiles of the adult peccary showed at least six dk, to the fractions: albumin, alpha-l, alpha-2, beta-l, beta-2, and gamtr 3. Globulin fractions of the peccary had different mobil mobilities from 1 4. The only sexual dimorphisms were associated with the t h e beta globuli globulin ratio. 5. Ingestion of colostrum in 1-day-old neonates was marked r by globulins. 6. The only significant difference between pregnant and tlactating la ~taL|tx~ tfem ~txtaJ~ ~ao ill ttt~ globulin ratio. Lactating females had higher concentrations concentratiq of alpha-2 globulin females.

INTRODUCTION Relativel' elatively few studies have dealt with physiology of the le collared peccary (Tayassu tajacu; Tayassuidae) (Sowls, ;owls, 1961; Low, 1970; Zervanos and Hadley, 1973; Zervanos ervanos, 1975; Sowls et al., 1976; Zervanos and and none have examined the composition D .aay, 1977) . . . . . . . . of serum proteins. Various I~rotein fractions of serum lical indices of metabolic have shown promise as clinical 'ouraine et al., 1982; Gay protein status in humans (Touraine and Touraine, 1982; Moskowitz :owitz et al., 1983). Recently, there has been a considerable amount of research directed toward develo eveloping similar indices for assessing dietary proteinI status of wild ungulates (LeResche et al., 1974; Franzmanr anzmann ann and LeResche, ~1 and Hoskinson, 1978; 1978; Seal et aL, 1978; Seal Warren et al., 1981, 1982). Inn domestic swine, a close taxonomic relative of the peccary, serum proteins have been shown to vary according to nutrition (Lecce and Matrone, 1960; Lecce, 1961; Lecce et al., 1962; Venum et al., 1970; Tumblest Fumbleson, leson 1972; Yen et al., 1982). The usefulness of.f protein profiles in animals also has been demonstr aonstrated for forensic identification (Wolfe, 1983)) and systematic classification (Johnson and Wicks, 1959). As part of a larger studyy investigating the physregulate, serum proteins iological life-cycle of this un rose gel electrophoresis, were characterized by agarose Using blood serum collect~ed from both wild and :aries, we examined procaptive Texas collared peccaries, tein profiles as influenced by, sex, reproductive condition, and age.

zed by agarose gel ly-active females in ands corresponding wine. io and the albumin: increase in gamma alpha aq-,lta globulin: ~bl, beta than non-pregnant no

were trapped in modified Clover deer traps from Zavala Coun Texas, using corn as bait. Animals County, Anir were sexed and classi classified as adult or juvenile using dental dent eruption patterns (Kirkpatrick and Sowls, 1962). Animals Animal were immobilized using ketamine hydrochloride administeered intramuscularly by blow-gun bh syringe (Lochmiller and Grant, Gn 1983) at a dose of 20 mg/kg body weight. Blood was collected for serum t -. . . . tube assemblies from the anterior vena cava using vacuum vacu pies were placed on ice (Lochmiller et al., 1984). Blood sample, isolated by centrifugation after collection and the serum was isolat (2000 g, 20 min) and frozen at -20°C until analyzed. Blood samples also were collected fror a group of captive ,~ted from animals using immobilization and ble~ bleeding techniques as described above for adults. Nursing young yo were bled from the orbital sinus (Lochmiller et al., 1984). 198 At 12-24hr, 21 days, and 42 days after birth collection ~ollections were made from pregnant, lactating and nonpregnant adult a females main~estible energy/kg dry tained on a high energy (3500 kcal dige weight), high protein (16~) pelleted ration provided ad libitum.

Protein chemistries Serum total protein was determined b~ythe Biuret method using the Technicon SMAC system (Te Technicon Inc., 1981). Separation of serum proteins was carrie~ ed out by agarose gel electrophoresis with a LKB-Produktc adukter 2117 Multiphor (LKB Produkter AB, Bromma, Sweden) Swede at a pH of 8.6, using Tris-barbiturate buffer. Serum sa samples (200/~ 1) were diluted with a 0.1~ bromophenol blue solution (100/~1) as the tracking dye. A 2/~1 sample of diluted diluter serum was applied to the gels. Electrophoresis was carried out at 20 V/cm for 40 min. Gels were fixed in picric acid, and stained in 0.5~ amido black and destained in 10~o acetic a~ acid. Relative percent of each protein was analyzed b) by scanning gels on a LKB-Produkter 2202 Ultroscan Laser densitometer with

M E T H O D S

Sample collections Blood was collected from 32 wild and 2

It males and females were ay analyses of variance with

ROBERT L. LOCHMILLER el al.

among pregnant, lacptivity also were tested ance with reproductive alyses were performed zedure of the Statistical il, 1979).

r'pical protein patterns At least six distinct protein bands were evident on ectrophoretograms of adult peccary sera (Fig. 1). ensitometric scanning revealed a consistent pattern )mposed of six distinct peaks which closely resemed expected levels of albumin, alpha-1 globulin, pha-2 globulin, beta-1 globulin, beta-2 globulin, ld gamma globulin, anodally to cathodally, re~ectively (Fig. 2), when compared to domestic swine ra. Thus, we have adopted this nomenclature for ,'signating the six major protein fractions in the )llared peccary profile. Serum from an adult female ral swine (Sus scrofa) was collected in Zavala ounty, Texas, and its proteins were separated elecophoretically for comparison of mobilities to the mcary (Figs 1 and 2). The peccary albumin fraction tended to migrate lly slightly anodal to the swine albumin (Fig. 1). onB lose visual inspection of electrophoretograms of the Close ins tult peccary revealed a number of faint bands which adult ere associated closely with alpha and gamma globwere in fractions. These bands normally were not deulin ctable by densitometric scanning. One of these faint tectable rods, which appeared consistently in the adult pecbands.

cary, was Ioca lbumin alpha-I globulin interzone, just alpha-1 globulin fraction (Fig. 1). A si] ocated in this interzone region was es ninent among 1-day-old neonates (Fig. -2 globulin fraction often separated into tt bands of nearly equal intensity, but n on the densitometer (Fig. 1). A narrow ,ing intensit.~ and clarity often appeared lal region of the gamma globulin fracti rod, when present, was subst stantially ce beta-2 fraction, so it was considered a g~ n (Fig. 1). This band was consi eek-old nurslings tFigs 2 very prominen g captive pregnant and and 3), and 1, ?aptive adult males infrelactating fema lacta nd in this region. Serum quently showe quen females rarely produced from wild adu a region. bands in the a banc possessed by two wildSerum prot Se ;ly characteristic of adult tght juvenil, caug atterns of 1-day-old neo)atterns. Typi, patt{ -week-old nurslings a,e nates, 3-week nate~ distinct peaks are disshown in Fi~ ric scanning among net)tingtuishable b Fig. 2). Alpha-1 gk)bulins nates ingesting natee defined fractions, desepa:arated into two clearly defin tectable by the densitometer. Alph pha-2 globulins and tecta the beta globulins are diffuse among neonates become very promi(Fig. 2). The gamma globulins be~ nent after ingesting colostrum. /A neonate which ~lostrum, as evidenced by ~arently did not receive colostrur appa stomach at necropsy, [had only a slight an empty e: fraction .just cathodic indication of a gamma globulin fra~ indic to the beta globulins (Fig. 3).

ADULT MALE

LACTATING ADULT

PREGNANT ADULT

-ap

PIG

-

b2-..

b,-I

a2~

alb-A

n B

Fig. I. Typical patterns obtaine adult male (G, H) collared pecc is shown (1). Alb = albumin; a~ =

m n

£

E

D

BI ...... r, It G

g female (D F) and aduli feral pig also ;lobulin: b e = beta-2

571

Serum proteins of the collared peccar

6--Wl

g

alb

i/1.

/ T W Y

Ilk

|1

ol2

g

a~

I;I 112

g

alb

|

Fig. 2. Densitometric tracings of electrophoretograms fron from a l-day-o esting colostrum, a 6-week-old nursling, and a pregnant collared peccary.'. Also sl shown for re msitometric tracing Symbols are aennea in rig.;.11. of an adult feral pig electrophoretogram. aymools S At 3 weeks of age, electrophoretic profiles began to resemble semble those of the adult. Unlike the neonate, the allpha- 1 fraction appeared as a single intensely stained band, and the alpha-2 globulin was more clearly visibh fible. The two beta globulin bands also were well delinq lineated as in the adult. A faint band in the anodal gamma mma region was barely visible at 3 weeks, but became came conspicuous by 6 weeks of age (Fig. 3). The onl, dy detectable changes in mobility during the nursing period of development were associated with the

1 -DAY

alpha- 1 globulins. Neonates had a double alpha-I band which consisted of a band sliightly anodal and one sslightly cathodal to the location locatic of the alpha-1 globr lobulin in all other age groups (F (Fig. 3).

Concentration o f serum proteins Re Relative percent and absolute cconcentrations of serun constituent ratios found serum proteins and selected constitl in wild adult male and female pecca )eccaries are presented

6 ~-WEEK

3-WEEK

ap

..... g

.......

b2

...... ii 2

..... a 1

mla m A

B

C

Fig. 3. Typical patterns obtained gel electrophoresis. (A) Electroph of 1-day-old neonates ingesting (

P

r

oral i ~

ILl H

--lib I

)roteins by agarose ~ctrophoretograms reek-old nurslings. ed in Fig. 1.

ROBERT L.

LOCHMILLER

el al.

dimorphisms in the globulin:albumin )umin:globulin ratio ly different between a B:A averaging of 0.82_+ 0.05 (SE) ad 0.90_+ 0.04 (SE),

pregnant and hales. Lactating females had a larger A onpregnant and pregnant females. Relati tions of alpha-2 globulin were significa 05) lower among nonpregnant (3.2 mong lactating (4.47'! o) females. Diffe solute concentration of alpha-2 globul d significance (P < 0.06). Nursing yo ~d protein patterns dislant protein fraction tinctly differen animals (Table 2). Relatal serum protein in rive and absol ations of many fiactions ild adult peccaries. Albumin was followed by remained rema dissi the adult thoughout the ~mma globulin (301}(,) in relative abundance. Within nursing nursi perio( 6 weeks of age). The e alpha globulins, the alpha-I fraction (57 717%) 1-daly-old neor t-like total serum protein mtributed the largest proportion of protein (Table levels level: [7.67 _+ C ] after ingestion of COldS. The beta-1 fraction (59-66°i,) was the most abun- trum. trum A neon of colostrum had only mt beta globulin (Table 2). However, the beta 4.00 g/dl total nlike the adult, gamma ~7%], obtained from coloslobulins [53.3 obulins did not separate clearly in most wild pec- glob~ 1 the largest protein fracry sera so values were not presented, trum absorpti( was reflected in a high Serum total protein levels of the two wild juvenile tion in the m Ltio (G:A) and a low A : G ales were 28% below adult concentrations. Relative gam] amma globuli accounted Ibr only 8.6'!. otein concentrations differed as well. Relative per(Tab 2). G a n (Table the colostrum deprived ntages of albumin, alpha-1 globulin, and beta of total t( serun obulin exceeded adult values as the gamma globulin neonate. is separated into two disactions of the juveniles was 401'o lower than relative N( Neonate alp (Fig. 2). These uaHu~ U~L~L~UJ~ L,y U ~ Hnsitometry OILUIXI~ [ult levels (Table 1); however, absolute concen- tinct bands det both before )ha-1 bands were present in neonates neo ations of these fractions were all higher among alph~ The anodal alpha- I a [ults. and after colostrum ingestion. Th~ adults. :ntrauon of 1.63 +0.37 Female reproductive state had some influence on band had a relative concentratk Tt cathodal alphalative and absolute distributions of protein (Table (SE)')?o, or 0.12 _+ 0.02 (SE) g/dl. The relative was more prominent, having a relative lb band t 2).. The alpha globulin:beta globulin ratio (A:B) or 0.19 + 0.02 ,ncentration of 2.43 + 0.23 (SE)'~,, (SE) (P' < 0.01) was the only significant difference between conc

Table 1. Comparisonof rrelative e l a t i v e and a n d aabsolute b s o l u t e aan mounts of protein fractions, alpha globulin:albuminratio, beta globulin:albuminr'. ratio, alpha globulin:beta globulin ratio, gamma globulin:albumin ratio, and alb albumin:globulin ratio among wild adult and juvenile collared peccaries collected t¥om southern Texas

ProteinI f r a c t i o n ( u n i t s )

WilJ adult m31es (~1 : I ( )

WilJ aJult females

Wild ]uvnnil~s

(~I : 1 ] )

(N = 2 )

Mean

'loan

0.16

7.23

'47.00

I . /')

3.54

0.12

~. 99

absolute (g/dl)

SE

. ,r

a5, :19

1.03

5b.~'%

1.28

0.0~

2.38

O. lb

O. 55

% ~15

0.23

g. 15

O.?u

0.52

O. 04

O. qT

0.02

0.42

0. Ol

~1.86

0.36

5.4?

0.15

5.10

),23

Albumin~, relative (%)

A l p h a - 22 globulin, relative (%)

!lea~

5,~0

7.55

A l p h a - 11 globulin, relative (%)

'SE

0.1g

Total pprotein ( g l d l )

absolute (gldl)

:;E

!.Y'

absolute (g/dl)

0.37

0.03

0.40

0.02

0.~5

0.01

T o t a l alpha globulin, relative (~)

11.80

0.81

II.85

0.20

13.25

9.9)

0. q )

absolute (g/dl)

T o t a l beta g l o b u l i n ,

relative

absolute (g/dl) globulin, relative (~) Gamma gl absolute (g/dl)

Alpha globulin/albumin ( r a t i o )

(~)

O.B9

0.07

0.88

0.03

0.69

12.47

0.~1

13.1j

0.~0

13.95

1.3>

0.9~

0.0~

O.9T

0.05

0.72

0.04

29.18

1.87

]0.91

I.~3

18.09

1.30

2.29

0. I.

2.33

0.17

0.93

0.19

0.25

0.01

0.25

O. Ol

0.2:~

O. Ol

Beta globulln/alt

).;!3

Alpha globulin/b(

3.o9

Gamma globulin/a

2.02

Albumin/globulin

3.02

Serum proteins of the collared pecca

573 beta globulin:albumin ratio, alpha g at birth, 3 weeks, and 6 weeks ot

te amounts of protein fractions, alpha gobuli~ albumin ratio, and albumin:globulin ratio amo nd among pregnant and lactating females whil

Captive females

Nursing young After eolostr um

3-weeks

ingestion

old

old

L~etating

(,S:3)

(N_:3)

(LI=6)

(N=3) Mean ~rotein (g/dl) ~, r e l a t i v e (%) absolute (gldl) I globulin, relative (¢) absolute (g/dl) 2 globulin, relative (%) absolute (g/dl) alpha globulin, relative (%)

absolute (g/dl)

Mean

S£

absolute (gldl) Alpha globulin/albumin 3~ta globulln/albumin

(ratio) (ratio)

SE

t~n

SE

0.35

7.67

0.35

6.00

0. I0

5.97

0.14

7.5B

0.17

7.35

1.57

51..3

2.58

53.50

1.62

"0,77

1.76

a5.32

1.79

2.62

2.37

0.05

3.08

0.1o

3.19

0. o2

3.08

0.09

3.34

0.25

6.10

,.07

0.37

10.77

1.07

11.13

O.q3

8.23

0.55

7.~J5

0.32

0.24

0.31

0.02

0.6"

0.05

0.66

0.01

0.62

0.05

0.58

0.05

1.90

2.30

0.75

3.97

0.18

3.43

0.28

,.47

0.3~

3.20

0.33

0.08

0.18

0.06

0.2"

0.01

0.20

0.02

0.3U

0.03

0.24

0.03

8.00

6.37

0.69

14.73

1.01

I,.57

0.41

12.70

0.57

11.05

0."3

0.32

0."9

0.07

0.88

0.05

0.87

0.02

0.95

0.06

0. B2

0.07

11.07

0.,9

10.50

0.65

9.23

0.5B

11.1o

0.90

0.66

0.03

0.62

0.02

0.7C

0.02

0.70

0.04

0.82

0.10

6.60

0.30

5.87

0.58

6."0

0.28

5.38

0.39

5.75

0.23 0.03

0.,0

0.02

0.35

0.03

0. q8

0.02

0.49

0.04

0.42

18.00

9.g0

0.85

17.70

0.55

16..0

o.91

15.85

0.17

15.60

0.72

16.87

0.89

0.72

0.76

0.07

1.05

9.04

0.98

0.03

1.18

0.02

1.1B

0.07

1.24

0.13

8.60

53.]0

0.98

17.13

3."2

15.90

2.75

31.62

1.93

31.73

1.28

27.55

2.3"

O.,A

".09

0.25

1.03

0.22

0.96

0.18

2.37

0.20

2.qI

0.14

2.01

0.15

absolute (g/dl)

Gamma globulin, relatlve (%)

Mean

SE

31.10

globulin, relative (%)

absolute ( g / d l )

1lean

(L~=-~)

".00

absolute (g/dl)

relative (%)

5£

tJo~-reproductlve

65.q0

globulin, relative (%)

Total beta slobulln,

Mean

6-~eeks

O. 12

0.21

0.03

0.29

0.01

0.27

0.01

0.27

0.01

0.32

0.03

0.2.

o. oi

0.28

0.32

0.0~

0.35

0.02

0.31

0.01

0.38

o.oi

0.39

0.03

0.37

o.01

Alpha globulin~beta globulin ( r a t i o ) O . , ,

0.64

0.04

0.83

0.06

0.89

0.05

o.73

O.OB

0.81

0.01

0.56

0.03

Gamma globulin/albumin ( r a t i o )

O, 13

1.72

0. II

0.3,

0.08

0.30

0.06

0.76

0.08

079

006

062

0.07

Albumln/globulin (ratlo)

1.89

o.,5

0.03

1.os

o.11

1.1,

o.o8

0.72

0.05

0.69

o.o,

0.82

0.05

(SE) g/dl. Total alpha-1 concentrat ncentration ration was half the level found in the adult peccary. cary. Total levels of beta globulin among neonates also so were well below characteristic adult. By 3 weeks of age substantia ~bstantial ~tial developmental changes became apparent (Table 2). Total protein concentration decreased by 2 2 ~ to a level of 6.00 + 0.10 (SE) g/dl. The gamma globulin fraction experienced a 70% reduction from neonata neonatal atal levels. Albumin increased to an absolute concentr )ncentration [3.08 +0.10 (SE) g/dl] resembling the adult ult peccary. Two alpha-1 bands were no longer detectable ctable and both relative and absolute amounts of this ais fraction had doubled since birth. A slight increase,~in alpha-2 globulin was also noticed when comparedd to neonates, but levels remained below the observed ed adult level. Also, the beta globulins had separated ted into two adult-like bands by 3 weeks of age. Concomit oncomitant to this separation was a 40~o increase in beta globulin over neonatal concentrations (absolute). )solute). However, there was little difference in the B:A [0.32 + 0.04 (SE)] and 3-week-old [( nurslings. Total beta globulin con weeks of age was similar to adult le

terns between 3-week and 6-week-old 6-we nurslings (Table 2). The most obvious differe, erence between adult and 6-week-old electrophoretogra rams was the extremely low gamma globulin concentration con, of the latter; 60% below adult levels. DISCUSSION The six serum protein bands ch~ characteristic of the adult collared peccary are in agrreement with the number reported for domestic ic swine (Kaneko, 1980). However, globulin mobilities differ slightly between these two species. The overall mean (N = 14) concentration of albumin was 3.17 + 0.08 (SE) g/dl in the captive females, which was in the th( reported range (1.80-3.30g/dl) for swine (Kaneko, 1980). Distribution and concentration of alpha globulins in the peccary are also apparently diffe~ different from swine. Usually, the alpha-2 globulin fraction frac is the most abundant alpha globulin in swine (1% (Miller et al., 1961; aeko, 1980); however, in ;ion predominates. Swine • overall alpha globulin ) be differences in distribeta globulins between

ROBERT L. LOCHMILLER e t al.

al., 1961; Kaneko, tion is the predomaeko, 1980), whereas undant in peccaries, + 0.09 (SE) g/dl of published range of o, 1980). sexual dimorphisms o has been reported al., 1961). Various searchers have elucidated the protein changes ac)mpanying gestation and lactation in swine (Foster al., 1950; F r i e d e l l e t a l . , 1951). F r i e d e l l e t a l . (1951) ported that alpha globulin concentrations remained irly constant during gestation, but were highly triable during lactation. Alpha-2 globulins tended increase during pregnancy and lactation in the male peccary. Friedell et al. (1951) also noted a :neral downward trend in albumin levels during ctation and a downward trend in gamma globulins 1ring gestation. These protein responses were not ~served in this study'. Several studies have documented serum protein enamics during the first few hours after birth in /ine (Rutqvist, 1958; Lecce and Matrone, 1960: Tiller et al., 1961; Asplund et al., 1962; Lecce et al., )62; Waddill et al., 1962: Pirtle and Deyoe, 1963; Masck 7asek and Holub, 1970). Developmental changes :curring in nursing swine appear similar to the occurrln ranges observed in nursing peccaries. Rutqvist chan 958) observed that the beta globulins derived from (1958 colostrum in swine were heterogeneous electromretically, and thus indistinctly separated from one phoreticall lother. This also appears to be a characteristic of another. :ta globulins in the peccary neonate (Fig. 3). beta Prior to nursing the serum protein concentration in newborn pigs averages 2.2 g /dl (Miller et al., 1961 ) to 2.4 g/dl (Lecce and Matrone.e, 1960). A newborn pecmd a much larger concencary deprived o f c o l o s t r u m had 10 g/dl (Table 2). Alpha-2 tration of serum protein, 4.00 globulin makes up more thann half halt • and an albumin about atein in newborn swine one-fifth of the serum protein before suckling (Lecce and Matrone, 1960: Miller et al., 1961; Pirtle and Deyoe, 1963). in the colostrumdeprived peccary neonate albumin was the most abundant fraction, contributin uting over 65'I o of total ~ere was no difference in serum protein. However, there in when compared to the absolute quantity of albumin suckling newborn peccary ('Table 2). Ited for 0.34g/dl (8.60!!~;) G a m m a globulins accounted of the serum proteins in the colostrum-deprived und no gamma globulin peccary. Rutqvist (1958) found present in the serum of pigs at at birth: however, Foster et al. (1950) noted a small amount mount, while Miller el al. (1961) reported 6.5'!~, and Pirtle and Deyoe (1963) 5.0!~i. Following the consumnption of colostrum and action increased to levels peccary g a m m a globulin fraction higher than those reported for nursing swine (Miller et al., 1961). No apparent differences in the beta globulin fraction were notedd between colostrum-fed and deprived peccaries. This; observation differs from nursing swine where beta globulin significant portion of the colostrt crease (Miller et al., 1961). Speer et al. (1959) have shown th

after birth i colostral-prote gamma globul the peccary. 6-week-old pe ported that ga in swine at al virtually no through 3 wee mins begins so does also in pe. pha globulin alpha 9ment in swi opm, elevations in t eleva peric )eriod from bi levels level: were 1o~ 9ment chang, opm(

)iminishing supplies ol cted in the extremely low g/dl) at 3 weeks of age in levels persisted in the • Miller el a/. (196t) ren levels reach a minimum s of age, indicating that mlin production occurs "oduction of serum albuin swine, and apparently ,"2). The amount of serum e constant through develtot in the peccary. Large action occurred over the :s of age (Table 2). Adult that postweanling develred.

Ackn

Acknowledgemet cial support provided by the Caesa Caesar Kleberg )gram in Wildlife Ecology, Depa )artment of ;heries Sciences, Texas A&M Univ, University and 1 title Association of America are gratefuly a{ We thank the staff" of the Chap)arrosa Ran Fexas for providing access to S. Heineman for providing wild collared pe technical assistance in the laboratory.

REFERENCES

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