ARCTIC
VOL. 38, NO. 1 (MARCH 1985) P. 6184
Selenium Concentrations in Forages
of a Northern Herbivore
GEORGE G. SHAWLand HAL W. REYNOLDS*
ABSTRACT. The importance of adequate selenium in diets of native wild herbivores
can only be inferred fromdata for beef cattle whereminimum
dietary concentrations range from
50 to 100 ppb. Concern about possible selenium deficiencies in wild herbivores
based
is on a few reports of symptoms in wildlife, a paucitydata
of on selenium in theirforages,and the idea that excessive atmosphericsulfur may increasethe incidence of selenium
deficiencies in herbivores. Concentrations of selenium insedges, Carex spp., and reedgrasses, calamagrosris spp., the main food plants of bison,
Bison bison, in northwestern Canada, varied from 9 to 800 ppb in samples collected at three lowland locations. However, approximately threequarters ofall the samples of plant species consumed by bison were dietarily deficient by the beef cattle standard.
Key words: Carex, calamagrostis, bison, selenium concentration
&UMfi. L’importance d’un quantit6adkquate de sklbium dans les rCgimes des herbivores sauvages locaux ne peut &re infCr6e que
d’apks les
donnh portant sur le Mtail bovin, dans lequel les concentrations minimales varient entre 50 et 100 ppb. La question d’insuffisance possible en
dlenium chez les herbivores sauvages fut lev& par rapport h quelques etudes des sympt8mes d’animaux sauvages, la p6nurie de donnh sur le
dlCnium dans leur rkgime et h la hypothhe selon laquelleun ex& de souffre dans I’atmosphZxe pourrait augmenter I’incidence d’insuffisance en
&lCnium chez les herbivores.Les concentrations de dlCniumdans les laiches(Carex spp.) et les gourbets( C h g r o s t i s spp.), les plantes alimentaires principales dubison (Bison bison) dans le nord-ouest du Canada, variaient entre
9 et 800 ppb dam les 6chantillons recueillis
h trois sites en terres basses.Cependant, environ les trois quartstous
de les 6chantillons tiese s w s de plantes consomm&s
par le bison furenttrouvh insuffisants au
niveau alimentaire en comparaison avecla norme pour le Mtail bovin.
Mots cMs: Carex, C h g r o s t i s , bison, concentration de SeICnium
Traduit pour le journal par Maurice Guibord.
INTRODUCTION
ported or demonstrated in other captive herbivores (Sauer and
A small amount of selenium (Se) is essential in the diet of
Zook, 1972; Brady et al., 1978; Kurkela, 1980). However,
warm-blooded vertebrates. Selenium forms part of the enzymeconclusive evidence that native free-ranging herbivores may
glutathioneperoxidase(GSH-Px)andhasotherimportant
suffer from Se deficiencies in suspected Se
lowareas is lacking
1976; Stadtman,
metabolicfunctions(WelshandSoares,
except in the case of some Oregon voles (Pond et al., 1982;
1977; Harr, 1978). Hoekstra (1974) explained the role of Se
Whitfield, 1978).
and the biochemistry of GSH-Px in the prevention of such Se
Seleniumdeficiencydiseasesofwildherbivoresinsusdeficiency
diseases
as
white
muscle
disease,
exudative
pected low Se areasmay not be reported because carcasses of
diathesis, and weak calf syndrome. The recommended minidead herbivores sufficiently freshfor the diagnosis ofSe defimum dietary concentrations of Se for beef cattle are 50-100
ciency symptoms would be difficult to find, and if found, the
ppb (Prior et al., 1982) and may apply also to wild herbivores tissues may be decayed or may have been consumed by other
such as bison.
animals. On the other hand, Se deficiency diseases may not ocDuring the last decade, several surveysof agricultural soils cur in wild herbivores for the following reasons. The Se conand cultivated forages in the boreal forest region of Canada
tent of cultivated foragesmay not be representative of uncultifound that a high proportion of forage samples was dietarily
vated plant species that are growing on soils with low Se condeficient in Seby the livestock standard (Levesque,1974a,b,c; tent or availability. Wild herbivores normally may encounter
Gupta and Winter, 1975; Redshaw et al., 1978; Winter and
uncultivated forage plants with moderate to high concentraGupta, 1979; Westra, 1982). Similarly, a large proportion of
tions of Se, thus obtaining sufficient dietary Se.And finally, it
forage samples in the northern U.S.A. was deficient (Kubota
ispossiblethat“native”wildherbivoreshaveadapted
et al., 1967). Based on these data, it is conceivable that there throughevolutiontodietslowinSe
in comparisontothe
may be a general Se deficiency in forages fed upon by wild
standards set for livestock.
herbivores in the North American boreal forest region. HowBecause the preceding hypotheses have not been supported
ever, if this is true, why are Se deficiency diseases not more
by empirical data, the question remains: how frequently and
frequently reported for wild herbivores?
under what conditions do wild herbivores suffer from Se defi(1971), Fleming et al. (1977), and
HebertandCowan
ciencies? The need to answer this question has been given imStoszek et al. (1978) have reported symptoms of either Se or
petus by the idea that increased atmospheric sulfur from the
vitamin E deficiencies in free-roaming mountain goats, wood- combustion of fossil fuels (acid rain)
may decrease the concenchucks, and antelope respectively, although the causes of the
tration of Se in vegetation over large areas ofNorth America
symptoms were not confirmed. Symptoms have also been re- (FrostandIngvoldstad,
1975). Hurd-Karrer (1935,1937),
‘Corporate Affairs, Environment Canada, 804 - 9 w 2 - 108 Street, Edmonton. Alberta, Canada T5K 2J5
2canadian Wildlife Service, lo00 - 9w2 - 108 Street, Edmonton, Alberta, Canada T5K 215
G.G. SHAW and H.W.REYNOLDS
62
Davies and Watkinson (1966), and Gissel-Nielsen (1973) have Average concentrations of Se for sedges, grasses, forbs, and
shown Se reductions in several crops to be associated with ex-other non-forbs were compared by t-tests (PI 0.05) for difcessive elemental S and SO4 in soils.
ferent wet meadow habitats. Transformationof raw datato the
Gissel-Nielsen (1973) reported a reductionin Se concentralogscalewasusedtoreducevariances.Fourteen,ten,and
tions in barley and red clover from soilSO4fertilization. Soils eleven plant species were analyzed from the Slave River lowwere fertilized to 12.5 and 125 ppm of sulfur. Fertilization
lands,NahanniRiverlowlandhabitats,andtheHay-Zama
with SO4 markedly reduced the Se concentration in clover at
lakes area respectively.
all soil selenite concentrations, even those low enough to cause
50 ppb. However,
initial clover Se concentrations of less than
RESULTS AND DISCUSSION
despite these results and other preliminary experiments (Shaw
Concentrations of Se in planttissues wereso variable thatno
andCocks,1982),muchresearchisneededtoestablish
comparisons between sedges, grasses, forbs, and other nonwhether SO2 or acid rain SO4will lead to lower Se concentraforbs were statistically significant (Tables1,2, 3). The coeffition in vegetation and subsequently to an increased incidence
cient of variation for duplicate determinations (Table 1) was
of Se deficiencies in wild herbivores.
l o % , indicating that analytical precision was good. Variability
Se concentraThe purpose of this report is to present data
on
shown for the concentration ofSe in plant tissues was probably
tions in several wild species of plants collected from three
due to differences between sites, between species and plant
of
riverlowlandhabitatswithintheborealforestregion
Salk spp.
parts, and between sampling dates. The twigs of
western Canada and to relate these data to the question of Se
were significantly lower in Se than the leaves on a log scale
deficiencydiseasesinbison.Thethreelocationsrepresent
(Ps0.05; Table 1). At Camp Meadow five of six species inareas of existing bison populations, of recently released wood
creased inSe concentrations from March to June, while at
bison (Bison bison athabascue),and of a proposed release site
Stan's Meadow seven of nine decreased in Se concentrations
for wood bison (Reynolds et al., 1982a).
during the same period (Table 1). Large variability
inplant
concentrations of Se within a single habitat has been reported
METHODS
before by Joblin and Pritchard (1981), where the concentration
Range quality and quantity were determined by systematic
ofSeinperennial
ryegrass varied abruptly and irregularly
plot sampling of vegetation along transects located in lowland across a pasture by as much as sevenfold.
et
sites of the Slave River, Northwest Territories (Reynolds
Bison feed primarily in wet meadows and tend to select plant
al., 1978),theLiard-SouthNahannirivers,N.W.T.(Reyspecies not quite proportionally to their seasonal availability
area, Alberta
nolds et al., 1980),andtheHay-Zamalakes
(Reynolds et al., 1978). The major species are sedges (Carex
(Reynolds et al., 1982b). The Se data reported were derived
spp.) and reedgrasses (Calamagrostis spp.) (Tables 1, 2, 3),
from vegetation samples composed from several plots in the
and together they can form up to87% of the plant biomass in
different wet meadow habitats where bison presently graze andsedge and wet meadows (Reynoldset al., 1978, 1982b). Bison
where reintroduced wood bison would be likely to graze. Se
(Salk spp.),yellowavens
willsometimesfeedonwillows
data from a few plant samples in drier sites are also reported.
(Geum aleppicum), slender wheat grass (Agropyron trachySe analyseswereperformed
by theSoiland Feed Testing
caulum), wire rush (Juncus bulticus), and horsetails (EquiseLaboratory of Alberta Agriculture in accordance with standardtum spp.) (Reynolds et al., 1978). When considering only the
above plant species as food items of bison, and the recomfluorometricprocedures(LevesqueandVendette,1971).
TABLE 1. Concentration (ppb) of selenium in tissues of 14 plants from two wet meadowhabitats in the Slave River lowlands, Northwest Territories, in 1975
~~~~
~~
Camp Meadow
Species
June
Carex
38 atherodes
21'
11 rostmta
10
Carex
Carex aquatilis
Scolochloa festucacea
10
syzigachne
Beckmannia
17
Calonrrgrostisspp. 15 37
295
A g q y r273
o n tmchycaulum
462
Hordeum jubatum
Pea spp.
Glyceria spp.
520
Juncuslmlticus
311
310
172 Geum aleppinun 6
Rumex maritimus 11
Mix spp. (leaves)
solix spp. (twigs)
'Duplicate determinations.
Stan's Meadow
March
20' 35
-
17
-
15
13
10
16
1447
438
-
-
6
10
-
32
34
16
1748
1683
483-
724
-
-
24
4181
14
12
15
44
20
1600
1560
450
18
19
670
21
41
22
9
37
5
22
76
8
35
-
-
34
332
36
21
45
23
49
40
20
144
136
23
45
-
-
-
-
20
22462
189
-
5
20
420
175
5
63
SELENIUM IN DIETS OF BISON
TABLE 2. Concentration @pb)of selenium in tissues of 10 p h t s in
Nahanni River lowland habitats in late June 1979
Habitat
Species
and site number'
Carex atherodes
Carex atherodes
Carex atherodes
Carex rostmta
Carex rostmta
Carex roam
Carex aquatilis
corur aquatilis
Carex aquatilis
Caknnagrostis cardensis
Caknnagrostis cardensis
Equisetum arvense
Equisetum pmtense
Equisetum pratense
Equisetum pratense
Equisetum palustre
Myrica gale (leaves)
salixmeadow
spp.
shrub
(leaves)
meadow
shrub
B
(leaves)
e d sp.
'Reynolds et al. (1980:Fig. 3).
Se
wet sedge meadow (3)
wet sedge meadow (5)
wet sedge meadow (5)
wet sedge meadow (5)
wet sedge meadow (3)
wet sedge meadow (5)
shrub meadow (1)
wet shrub bog (7)
wet shrub bog (7)
wet sedge meadow (5)
wet sedge meadow (3)
wet sedge meadow (5)
wet sedge meadow (5)
aspen sprue forest (4)
sjmlce. forest (2)
shrub meadow (1)
wet shrub bog (7)
39
250
640
400
635
31
220
64
67
64
800
40
670
3875
83
524
79
23
(1)
(1)
We thank the Soil and Feed Testing Laboratory of Alberta Agriculture for doing the !k analyses.
Sedge meadow Wet sedge-grass
meadow, Site H1
Site E'
Carex atherodes
Carex atherodes (tops)
(midstems)
(stem bottoms)
Carex aquatilis
Carex aquatilis (tops)
(midstems)
(stem bottoms)
Carex sp.
Eleocharis sp. (a)
Eleocharis sp. (b)
Scolochloa festucacea
Calamagrostis inexpansa
Beckmannia syzigachne
Typha latifolia
Alisma SD.
Bidens cernua
ACKNOWLEDGEMENTS
34
TABLE 3. Concentration (ppb) of selenium in tissues of 1 1
plants in
sedge meadow and wet sedge-grass meadow of the Hay-Zama lakes
area of northwestern Alberta in August 1981
Species
Because bison move frequently between meadows while grazing they are likely to consume daily some plants with concenet al.
trations of Se above 100 ppb.Moreover,Reynolds
(1982a)reportedbisontohaverelativelygreaterdigestive
of beef cattle. If this digestive superioricapabilities than those
ty applies to Se, the overall diet of bison may be adequate to
prevent Se deficiency diseases.
Theconcentrationsof Se reportedhereforseveralplant
speciesgrowinginthesamelocationswerequitevariable.
This suggests that considerable variabilityof Se concentration
in diets of wide-ranging herbivores may be expected, even in
areas where their main food items may be Sedeficient. This
should be confirmed by studies conducted on ranges of other
herbivores such as antelope and mountain goats, for which
E deficiencies have been reported
symptoms of Se and vitamin
previously.
30
60
25
32
9 102
20
9
10
BRADY,P.S.,BRADY,J.L.,
WHElTER, P.A.,ULLREY,D.E.,and
FAY, L.D. 1978. The effect of dietary selenium and VitaminE on bioamong white-tailed deer
chemicalparametersandsurvivalofyoung
(odocoileus virginianw).Journal of Nutrition 108: 1439-1448.
DAVIES, E.B., and WATKINSON, J.H. 1966. Uptake of native and applied
selenium by pasture species.I. Effects of sulphate and of soil type
on u p
take by clover. New Zealand Journal of Agricultural Research
9:641-652.
FLEMING, W .J., HASCHEK, W .M., GUTENMANN, W.H., CASLICK,
J.W., and LISK, D.J. 1977. Selenium and white muscle disease in
woodchucks. Journal of Wildlife Diseases13:265-268.
FROST, D.V., and INGVOLDSTAD, D. 1975. Ecological aspects of seleniumandtelluriuminhumanandanimalhealth.ChemicaScripta
81:%-107.
-
19
-
-
-
REFERENCES
34
'Reynolds et al. (1982b:Fia. 3).
mended level of 100 ppb
Se for beef cattle, 18 of 25 plant
samplesfromtheSlaveRiverlowlands(Table
l), 8 of17
samples from the Nahanni River lowlands (Table
2), and all of
the samples from the Hay-Zama lakes region (Table 3) were
dietarily deficient in Se. Three-quarters of all the samples of
plants known as food items for bison were deficient by the
beef-cattle standard.
However,speciessuchasslenderwheatgrass,wirerush,
horsetail, yellow avens, willow, and sedges sometimes have
concentrations of Se above 100 ppb (Tables
1, 2). A steady
diet of plants with 3900 ppb (horsetail - Table 2) would be
just subtoxic (Harr, 1978) but improbable, because such high
concentrations are not common among the samples analyzed.
GISSEL-NIELSEN, G. 1973. Uptake and distribution of added selenite and
selenate by barley andred clover as influenced by sulphur. Journal of the
Science of Food and Agriculture 24549-655.
GUPTA,U.C.,andWINTER,K.A.
1975. Seleniumcontentofsoilsand
crops and the effects of lime and sulphur on plant selenium. Canadian
Journal of Soil Science55: 161-166.
HARR, J.R. 1978. Biological effects of selenium. Chap.18 in: Oehme, F.W.
(4.). Toxicity of Heavy Metals in the Environment. New York: Dekker.
393426.
HEBERT,D.M., and COWAN,I.M. 1971. Whitemusclediseaseinthe
mountain goat. Journal Wildlife Management 34(4):752-756.
HOEKSTRA,W.G. 1974. Biochemicalroleofselenium.
In: Hoekstra,
W.G., Suttle, J. W., Ganther, H.E., and Mertz,
W, (eds.). Trace Element
Metabolism in Animals-2. Baltimore: University Park Press. 61-77.
HURD-KARRER, A.M. 1935. Factors affecting the absorption of selenium
from soils by plants. Journal of Agricultural Research 50:413427.
-.
1937. Selenium absorptionby crop plants as related to their sulphur
requirements. Journal of Agricultural Research54501-608.
JOBLIN, K.N., and PRITCHARD, M.W. 1981. Selenium in a ryegrasspasture. Proceedings, New Zealand Workshop on TraceElementsinNew
Zealand, University of Otago, Dunedin, New Zealand. 93-98.
KUBOTA, J., ALLAWAY,W.H.,CARTER,D.L.,CARY,E.E.,and
LAZAR, V.A. 1%7. Selenium in crops in the United States in relationto
selenium-responsivediseasesinanimals.JournalofAgriculturalFood
Chemistry 15(3):448453.
KURKELA, P. 1980. Green plant feeding of reindeer with reference to selenium. In: Reimers, E., Gaare, E., and Skjenneberg,
S.(eds.). Proceedings,
Second International Reindeedcaribou Symposium. Roros, Norway. Part
A: 207-212.
64
LEVESQUE,M. 1974a. SeleniumdistributioninCanadiansoilprofiles.
Canadian Journal of Soil Science W63-68.
-.
1974b. Some aspects of selenium relationships in Eastern Canada
soils and plants. Canadian Journal of Soil Science 54:205-214.
-.
1974c. Relationship of sulphur and selenium in some Canadian soil
profiles. Canadian Journal of Soil Science 54:333-335.
-and VENDETTE,E.D. 1971. Seleniumdeterminationin soil and
plant materials. Canadian Journal of Soil Science 51:85-93.
FOND,F.R.,TRIPP,M.J.,WU,A.S.H.,WHANGER,P.D.,
and OLDFIELD, J.E. 1982. Comparative effects of selenium deficiency on male
In: Gawthorne,J.M.,
reproductioninseveralmammalianspecies.
Howell, J.McC., and White, C.L. (eds.). Trace Element Metabolism in
Man and Animals. New York: Springer-Verlag. 365-367.
PRIOR,M.G.,ANGLE, R., GOATCHER,W.B.,GRAVELAND,D.N.,
LAVERTY, D.H., REGIER, H., SHARMA, R.M., and WESTRA, R.
1982. Theeffectsofenvironmentalsulphuronfarmanimals:
A pilot
study. Roceedings, AcidFormingemissionsinAlberta
and their
ecologicaleffects.SymposiumWorkshop,AlbertaDepartmentofEnvironment, Edmonton, Alberta, Canada. 551-570.
REDSHAW, E.S., MARTIN, P.J., and LAVERTY, D.H. 1978. Iron, manganese, copper, zinc and selenium concentrations in Alberta grains and
roughages. Canadian Journal of Animal Science 5833-588.
REYNOLDS,H.W.,GLAHOLT,R.D.,andHAWLEY,A.W.L.
1982a.
Bison. Chap. 49 in: Chapman, J.A., and Feldhamer, G.A. (eds.). Wild
M a m m a l s of North America. Baltimore, MD: Johns Hopkins University
Press. 972-1007.
REYNOLDS, H.W., HANSEN, R.M.,and PEDEN, D.G. 1978.Diets of the
Slave River lowland bison herd, Northwest Temtories, Canada. Journal
of Wildlife Management 42(3):581-590.
REYNOLDS, H.W., LYNCH, G.M., and LAJEUNESSE, B.L. 1982b.Unpublished ms. Range Assessment of the Hay-Zama Lakes
area, Alberta as
habitat for wood bison and a proposal for their re-introduction. Canadian
Wildlife Service, 1000-9942-108St., Edmonton,Alberta,CanadaTSK
215. 49 p.
REYNOLDS,H.W.,McGILLIS,
J.R., and GLAHOLT,R.D. 1980. Unpublished ms. RangeassessmentoftheLiard-SouthNahannirivers
region,Northwest Temtories as habitatforwoodbison.Canadian
WildlifeService, 1000-9942-108 St., Edmonton,Alberta,CanadaT5K
215. 39 p.
SAUER, R.M., and ZOOK, B.C.1972.Selenium-Vitamin Edeficiency at the
National Zoological Park. Journal of Zoo Animal Medicine 33-36.
SHAW, G.G., and COCKS, F.L. 1982.The effect ofSO2on food quality for
wild herbivores. Proceedings, Acid Forming Emissions in Alberta and
their Ecological Effects. SymposiumNorkshop, Alberta Department of
Environment, Edmonton, Alberta, Canada. 571-587.
STADTMAN,T.C. 1977. Biologicalfunctionofselenium.NutritionReviews 35(7):161-166.
STOSZEK, M.J., KESSLER, W.B., and WILLMES, H.1978.Trace mineral content of antelope tissues. Proceedings, Eighth Pronghorn Antelope
Workshop, Jasper, Alberta, Canada. 156-161.
WELSH, S.O., and SOARES, J.H., Jr. 1976.The protective effect of vitamin
E and selenium against methyl mercury toxicity in the Japanese
quail.
Nutrition Report International 13(1):43-51.
WESTRA, R. 1982.Sulfur and other mineral concentrations in feedstuffsfed
to livestock in various regions of Alberta. Proceedings, Acid Forming
Emissions in Alberta and their Ecological Effects. SymposiumNorkshop,
Alberta
Department
of
Environment,
Edmonton,
Alberta,
Canada.
589-607.
WHITFIELD, S.C. 1978.Some possible roles for selenium in the reproduc(Micronr~canicaudus) M.Sc.
tive physiology of the male gray-tailed vole
thesis, Oregon State University, Corvallis, OR, U.S.A. 95 p.
WINTER,K.A.,andGUPTA,U.C.
1979. Seleniumcontent of forages
growninNovaScotia,NewBrunswickandNewfoundland.Canadian
1 1.
Journal of Animal Science 59: 107-1
G.G. SHAW and H.W. REYNOLDS