Upload
mohamed-ali
View
11
Download
0
Embed Size (px)
DESCRIPTION
Geochemistry and the Mineral Nutrition of Agricultural Livestock and Wildlife
Citation preview
7/21/2019 Geochemistry and the Mineral Nutrition of Agricultural Livestock and Wildlife
http://slidepdf.com/reader/full/geochemistry-and-the-mineral-nutrition-of-agricultural-livestock-and-wildlife 1/12
Geochemistry and the mineral nutrition of agricultural
livestock and wildlife
Iain Thornton*
Environmental Geochemistry Research Group, T H Huxley School of Environment, Earth Sciences and Engineering,
Imperial College of Science, Technology and Medicine, London, UK
Abstract
Relations between the geochemical nature of soils and their parent materials and the occurrence of nutritional defi-ciencies and excesses in grazing livestock have been documented since the 1960s and earlier, with notable work in
Australia, New Zealand, United Kingdom, Ireland, USA and the Soviet Union. Studies at Imperial College London,
into the development of regional geochemical mapping techniques as a means of delineating nutritional problems
commenced in the 1960s and have continued to the present. Research has mainly focussed on deficiencies of Cu, Co and
Se in cattle and sheep, and the role of Mo and S in the soil/plant system in both clinical and sub-clinical hypocuprosis in
cattle. Exposure to heavy metals, in particular Pb, Zn, Cd and the metalloid As have also received attention in areas
contaminated by past mining and smelting activities. Soil and plant factors influencing the dietary supply of both essential
trace elements and toxic metals have been studied, including their speciation and bioavailability. Soil ingestion has been
recognised as an important exposure pathway of heavy metal contamination to grazing cattle, and as an antagonist of Cu
supply and a source of dietary Co in sheep. Relations between soil geochemistry and the mineral status of wildlife species,
in particular impala and black rhinoceros, have been established in Kenya and recent work, presented elsewhere in this
Symposium, has concerned the supply of nutrients to the Roosevelt sable in Kenya. The compilation of regional andnational multi-purpose geochemical atlases, based on the systematic sampling of soils or stream sediments, is now
recognised as a priority in many countries of the developed and developing world. This paper explores the opportunities
for future research into the application of geochemical maps for the optimisation of land use, efficient livestock produc-
tion, and improving conservation of wildlife.# 2002 Elsevier Science Ltd. All rights reserved.
1. Introduction
Mills (1979) reviewed the nature, pathological
response, metabolic defects, aetiology and detection of
trace element deficiency and excess in animals. He
emphasized the complexity of soil–plant–animal rela-tions with respect to trace element supply and noted
that the application of geochemical approaches to the
initial recognition of areas associated with a high risk
of anomalies in trace element supply to animals had
considerable potential value. Such an approach had
initially been proposed by Webb (1964), whose concept
of multi-purpose geochemical mapping was to instigate
wide-ranging studies in future years.
In a presentation entitled ‘‘Environmental geochemistry
and health in the 1990s: a global perspective’’ at the 2nd
International Symposium on Environmental Geochemistry
in Uppsala in 1991, Thornton (1993) reviewed the general
area of multi-purpose geochemical surveys and their
applications to agriculture and wildlife nutrition andmade some predictions as to priorities for future
research. The present paper draws attention to some of
the key features of early research, details progress over
the past 10 a and examines the outcome of these previous
predictions.
Studies have continued to focus on the same groups
of chemical elements relating to deficiencies of Cu, Co,
Se, etc., and excesses of Mo, Cd, Zn Pb and As
(Thornton and Webb, 1979; Thornton, 1983; Leech and
Thornton, 1987). The environmental geochemist has
continued to provide ongoing sources of information on
(i) background/baseline concentrations of elements in
0883-2927/02/$ - see front matter # 2002 Elsevier Science Ltd. All rights reserved.
P I I : S 0 8 8 3 - 2 9 2 7 ( 0 2 ) 0 0 0 7 9 - 3
Applied Geochemistry 17 (2002) 1017–1028
www.elsevier.com/locate/apgeochem
* Tel.: +44-171-594-3690; fax: +44-171-594-6408.
7/21/2019 Geochemistry and the Mineral Nutrition of Agricultural Livestock and Wildlife
http://slidepdf.com/reader/full/geochemistry-and-the-mineral-nutrition-of-agricultural-livestock-and-wildlife 2/12
rocks, soils and plants; (ii) spacial information on ele-
ment distribution in the form of geochemical maps and
atlases; (iii) the development of analytical techniques,
particularly for the characterisation of chemical and
mineral forms of elements in soils and (iv) arising from
the above, the provision of geochemical data required to
understand the processes leading to the solubility, bio-availability, and pathways of nutritionally essential trace
elements and potentially toxic metals in the rock–soil–
plant–animal system. Such information continues to be of
value in understanding the sources and supply/exposure of
elements to both domestic livestock and wildlife species.
In recent years, data handling, interpretation and
application has been assisted by the development of
Geographical Information Systems (GIS) and by the
development of risk assessment strategies used as tools
for the prediction of element deficiencies and excesses
leading to animal ill-health.
2. Agricultural livestock
Research in the 1960s and earlier into applications of
geochemistry to agriculture was mainly undertaken in
Australia, New Zealand, United Kingdom, Ireland,
USA and the Soviet Union. In the 1960s, 70s and 80s
much emphasis was placed on the nutritional needs of
domestic livestock. In developed countries, intensive
farming methods aimed to realize the genetic potential
of different breeds of cattle and sheep for meat, dairy
and wool production.
Within this period, geochemical maps, based on a
systematic sampling and analysis of stream sediments
and soils, were used to delineate areas of potential
nutrient deficiency and metal excess. Their main appli-
cation was to indicate areas or regions within which
more detailed study of soil and pasture herbage could
then be concentrated. For example, early studies in
Counties Whicklow and Carlow, Eire, indicated a
correlation between the Co content of stream sediments
and the occurrence and severity of pine in sheep and
cattle on soils derived from granite (Webb, 1964). Geo-
chemical maps showed similar low Co patterns (<10
mg/g) on granites in SW England on which pine hadbeen recognized since the 1930s (Patterson, 1938), while
moderately low patterns (10–15 mg/g) on the Culm
Measures in Devon and on Triassic drift in Denbigh-
shire were related to soil and pasture of low Co content
on which unthriftiness and pine were found in sheep
(Thornton and Webb, 1970).
One of the best examples of the application of geochem-
istry to agriculture is shown by the combined geochemical
map for Mo and Cu in England and Wales (Fig. 1) (Webb
et al., 1978) which highlights the considerable extent of high
Mo and low Cu land which is thought to exceed 4105 ha.
The main patterns of elevated Mo concentrations are
found in areas underlain by marine black shales
(Thomson et al., 1972). This elevated Mo in soil under
certain conditions is reflected in raised Mo levels in
pasture herbage which causes a conditioned Cu defi-
ciency in grazing cattle. As indicated in Fig. 2, the wide
distribution of low blood Cu values (hyopocupraemia) in
cattle is mainly located in areas with molybdeniferous soils,in which over 1700 herds were found to be Cu deficient
(Leech et al., 1982). Molybdenum accumulation in saltings
(recent marine alluvium) reclaimed some 400 a ago for
agricultural use is thought to be due to the fixation of Mo
by soil sesquioxides and organic matter. Extensive coastal
areas in Norfolk, Suffolk and Essex are affected as shown
by the geochemical map. Vegetation growing on these soils,
withpH values ranging from 7.0 to 8.0, was found to have a
Cu: Mo ratio below 4:1 and to be potentially toxic to
ruminants, depending on dietary SO4 (Smart, 1991).
As indicated by Thornton (1993), similar marine
black shales and marine/estuarine alluvium are found inmany parts of the world, though, with the exception of
North America, have received little attention as yet. It is
confidently predicted that as extensive systems of live-
stock farming are forced to become more intensive due
to population pressure worldwide, natural limitations to
food production of this type will become of increasing
importance. It is surprising that over the past 9 a little
further attention has been given to this emerging problem.
Perhaps in Europe one reason may be that less emphasis
is now placed on intensive livestock rearing, due to sur-
pluses of dairy products and meat arising as a result of
the EU Common Agricultural Policy. However, less
intensive production with reduced supplementary feed
has meant that grazing livestock are now more dependent
on grass as their main source of nutrition and thus
become dependent on the mineral status of the farm and
its soils. It is no doubt true that research into trace element
deficiencies is no longer fashionable in Europe and has
received little funding since the early 1980s. Nonetheless,
in the future, a less intensive and more sustainable system
of livestock farming will predispose towards nutritional
deficiencies inherent in the land.
The disease swayback, known as Lamkrius in South
Africa, Kipsiepsiep in Kenya and enzootic ataxia in
several other countries, including the former USSR, is anervous disorder of lambs. It is characterized by lack of
co-ordination of movement and paralysis of the back
legs, and has been long recognized in many parts of the
world (Underwood, 1966). It can be prevented by giving
Cu supplementation halfway through pregnancy. This
disease has been studied by a large number of animal
nutritionists, veterinary scientists, biochemists and
pathologists. However, an actual cause has been difficult
to find and a Cu-deficient diet would not seem to be the
simple solution.
Research in New Zealand clearly showed that farm
animals accidentally ingest appreciable amounts of soil
1018 I. Thornton / Applied Geochemistry 17 (2002) 1017–1028
7/21/2019 Geochemistry and the Mineral Nutrition of Agricultural Livestock and Wildlife
http://slidepdf.com/reader/full/geochemistry-and-the-mineral-nutrition-of-agricultural-livestock-and-wildlife 3/12
(Healy, 1968). Using Ti as a stable indicator of soil
ingestion, subsequent studies in the UK showed that
between 2 and 20% of the dry matter intake of grazing
cattle was soil, and up to 30 or 40% in sheep as they
graze closer to the ground (Thornton, 1974). This led to
an examination of the actual ingredients in soil that may
influence animal metabolism and nutrition.
Collaborative research with Professor Neville Suttle
of the Moredon Institute, found that 10% soil in the
diet of sheep reduced Cu absorption and utilisation by
Fig. 1. Combined element map for empirical Molybdenum (red), increasing colour contribution with increasing element concentration
and percentile Copper (blue), increasing colour contribution with decreasing element concentration.
I. Thornton / Applied Geochemistry 17 (2002) 1017–1028 1019
7/21/2019 Geochemistry and the Mineral Nutrition of Agricultural Livestock and Wildlife
http://slidepdf.com/reader/full/geochemistry-and-the-mineral-nutrition-of-agricultural-livestock-and-wildlife 4/12
as much as 50%, and that as little as 2% ingestion sig-
nificantly reduced Cu use. The mechanism for this was
not immediately clear and it was not known whether the
soil particles occluded Cu in the diet, whether they sup-
plied Mo, or whether the effect was due to some as yet
unknown process (Suttle et al., 1975).
In some simple laboratory experiments, soil was sha-
ken with rumen liquor in a test tube. The large amounts
of Fe coming into solution from this soil gave a valuable
clue. This work was repeated on a larger scale using
artificial rumen systems at the Moredon Institute. The
Fe released from soil is a powerful Cu antagonist, and it
was proposed that accidentally ingested soil provides a
source of large amounts of Fe in the rumen of sheep.
This Fe interacts with dietary Cu and reduces the utili-
sation of Cu. Suttle et al (1984) strongly suggested that
the soilCu antagonism was dependent on soil Fe, and
Underwood and Suttle (1999) present data indicating
the magnitude of the Fe effect. It would seem thus that
this mechanism could make a significant contribution to
the aetiology of swayback (Thornton, 1983), though this
requires further investigation. In the UK, swayback is
found in years with warm winters and very little snow
cover. In warm weather, sheep are left to fend for
Fig. 2.
1020 I. Thornton / Applied Geochemistry 17 (2002) 1017–1028
7/21/2019 Geochemistry and the Mineral Nutrition of Agricultural Livestock and Wildlife
http://slidepdf.com/reader/full/geochemistry-and-the-mineral-nutrition-of-agricultural-livestock-and-wildlife 5/12
themselves on poor pasture with little or no extra feed.
This is when soil ingestion is at its highest and when
supplementary Cu is needed by the pregnant ewe
(Thornton, 1983).
Soil may also contribute to the supply of nutritionally
essential elements for the grazing animal under UK
conditions and, in particular, the trace element Co.Cobalt is an essential constituent and, indeed, comprises
some 4% of vitamin B12 the anti-pernicious anaemia
factor of liver which is produced by the synthesizing
abilities of the ruminants symbiotic gastro-intestinal
flora (Underwood, 1962). Nutritional deficiencies of Co
result in severe emaciation and listlessness. These may
lead to death in both cattle and sheep; milder forms are
usually described as ill thrift and respond rapidly to Co
supplementation (Underwood, 1962). Cobalt deficiency
in sheep has been recorded in several areas in the UK
underlain by arenaceous course-grained soil parent
materials and by acid igneous rocks which contain onlyvery small amounts of Co. It is normally treated by the
direct oral administration of ‘‘cobalt bullets’’ to the
animal or by the treatment of deficient pastures with Co
SO4. As there is 100 to 1000 times more Co in soil than
in pasture herbage, potentially, a little ingested soil
could provide the dietary requirements of the animal,
irrespective of how much Co is present in plants. Fur-
ther collaborative studies between Imperial College and
the Moredun Institute, showed that not only was Co
extracted from soil in the sheep’s rumen, but it was also
synthesized into Vitamin B12 as required by the sheep
(Brebner et al., 1987). The diagnostic practice of MAFF
for the past 20 or more years had been based on the
analysis of pasture herbage for Co. On the evidence of
this research, it was suggested that this may not have
been appropriate as sheep directly ingest appreciable
quantities of Co in soil.
One of the possible limitations of much of the
research into relations between geochemistry and live-
stock nutrition has been that the majority of studies have
been based on the determination of ‘‘total’’ rather than
‘‘bioavailable’’ concentrations of mineral elements in the
soil. The use of chemical extractants such as EDTA and
ammonium acetate to estimate plant available levels of
trace elements have met with limited success particularlywhen comparisons are made between contrasting soil
types with large variations in pH or organic matter
content. Indeed it has been suggested that ‘‘total’’ con-
tent in combination with soil pH may well be a useful
estimate of bioavailability and may be used to predict
levels of trace elements in the soil solution (Rieuwerts et
al., 1998).
The Soil Geochemical Atlas of England and Wales
(McGrath and Loveland, 1992) provides maps showing
the distribution of both ‘‘total’’ and EDTA—extractable
trace elements on a sub-regional scale, though limited to
a sampling density of one per 25 km2.
With the research focus and funding moving from the
recognition and remediation of trace element defi-
ciencies, the move over the past decade has been
towards environmental protection with the emphasis on
human health and the ecosystem. An example of this
new perspective is provided by recent research colla-
boration between the Veterinary Laboratories Agencyand Imperial College which has focussed on the possible
influence of geochemical factors on the susceptibility of
cattle to infection by bovine tuberculosis from badgers
which is of increasing prevalence in the UK at this time.
Research has aimed to predict areas where deficiency of
Cu in the diet results from excess soil Mo whose avail-
ability and uptake into pasture is influenced by soil pH
and other soil factors. The geochemical maps for Mo
and Cu together with information on soil pH and S has
been introduced into a GIS system to predict possible
nutritional stress and increased likelihood of suscept-
ibility to infection. (Drs. P.A. Durr and M. Ramsay,1999, pers. commun.). This work falls within a broader
programme funded by the Ministry of Agriculture,
Fisheries and Food (Phillips et al., 2000).
In April 2000, Part IIA of the Environmental Protection
Act came into force in the United Kingdom—com-
monly termed the ‘‘Contaminated Land Regime’’. This
introduced the concept of ‘‘pollutant linkage’’ in which
meat and dairy products from the grazing animal can be
seen as part of the pathway from contaminated soil to
the human receptor. Heavy metal contamination of
agricultural soils arising from past mining and smelting
activities provides an appropriate example. On such
land, it is accepted that the principle threat to livestock
is directly ingested soil and vulnerable animals are those
consuming soil while grazing or foraging on con-
taminated land (Underwood and Suttle, 1999). Lead
mining in the Southern Pennines over several hundred
years to around 1900 has resulted in contamination of
around 250 km2 of land, with concentrations ranging
from several hundred to several thousand mg/kg Pb and
as much as 1% or more in and around mineral wastes
(Colbourn and Thornton, 1978). Rarely, cattle are
affected by Pb, usually through straying onto waste tips
and eating brightly coloured galena (PbS). A survey of
grazing cattle on 11 farms with soils ranging from 150 to4000 mg/kg Pb showed blood Pb levels to broadly
reflect amounts of Pb in the soil, as a result of ingesting
contaminated soil and pasture (Fig. 3). Between 40 and
70% of the Pb intake was shown to be as a result of
directly ingested soil (Thornton and Kinniburgh, 1978;
Thornton, 1983). This study also showed a wide range
of blood Pb levels encountered at any one time within a
single herd, probably reflecting the different rates of
absorption, metabolism and excretion between individ-
ual animals, coupled with different grazing habits of
individual cattle. It has been suggested that the greater
response in blood Pb to soil Pb recorded in the early
I. Thornton / Applied Geochemistry 17 (2002) 1017–1028 1021
7/21/2019 Geochemistry and the Mineral Nutrition of Agricultural Livestock and Wildlife
http://slidepdf.com/reader/full/geochemistry-and-the-mineral-nutrition-of-agricultural-livestock-and-wildlife 6/12
part of the grazing season compared to the late season
may be due to the initial metabolic response of the animal
to Pb exposure and changes in the rate of Pb metabolism
resulting from continuing exposure, a phenomenon also
encountered in the human (D. Barltrop, pers. commun.,1983). Although Suttle et al. (1991) have clearly shown
that the majority of Pb ingested in soil is excreted in
faeces, it is evident that a small proportion is absorbed—
an important factor on heavily contaminated land. Later
studies showed that in this area Pb contaminated soils
also contained appreciable amounts of F (some exceeding
1% F) and that soil ingestion was a major pathway of
soil F into grazing cattle (Geeson et al., 1998). Indeed, it
is recognised that ingestion of F-rich soil on overgrazed
pastures can significantly contribute to F intakes
(Underwood and Suttle, 1999). However, neither exposure
to Pb or F would seem to result in losses in production
or in the contamination of products offered for sale,
including milk. In this respect, Oelschlager et al. (1970)
had previously shown that ingested soil F was poorly
absorbed by cattle.
Another example is provided by the metalloid As.Extensive contamination of agricultural soils in SW
England has been documented by Abrahams and
Thornton (1987). Within the Hayle/Camborne area the
concentrations of As and Cu found in surface (0–15 cm)
soils across the Cornish Peninsula ranged up to 730 ppm
As and 560 ppm Cu, confirming the widespread surface
contamination of the area during the course of mining
and smelting operations spanning several centuries, with
around 700 km2 of agricultural land in Cornwall and
Devon affected to varying degrees (Abrahams and
Thornton, 1987). The agricultural significance of these
findings was, in part, assessed by determining the uptake
Fig. 3.
1022 I. Thornton / Applied Geochemistry 17 (2002) 1017–1028
7/21/2019 Geochemistry and the Mineral Nutrition of Agricultural Livestock and Wildlife
http://slidepdf.com/reader/full/geochemistry-and-the-mineral-nutrition-of-agricultural-livestock-and-wildlife 7/12
of As and Cu into pasture herbage and the intake of
these elements by grazing cattle on 11 farms in the
region, selected on the basis of the As content of their
soils. Neither As nor Cu are readily taken up and
translocated to the aerial parts of grasses, and it has
been clearly shown that accidentally ingested soil com-
prises a major exposure route to the grazing animal(Table 1; Thornton and Abrahams, 1983). Further
studies showed that the amount of soil ingested (calcu-
lated on the basis of the Ti content of faeces) varied
appreciably between seasons of the year and in this
study ranged from 0.2 to nearly 18% of the animal’s dry
matter intake (Abrahams and Thornton, 1994). In the
spring when there is little grass cover, this could account
for around 90% of the As and 30–60% of the Cu intake
of grazing cattle. Even though some of the animals
would seem to ingest in excess of 50 mg of As per day,
no adverse effects on the health or production of cattle
have as yet been reported in this area. Anecdotally, ithas been suggested that local cattle and sheep have
developed some resistance to biological exposure to
these two potentially toxic elements. Scanning electron
spectroscopy and X-ray analysis has shown that much
of the As in surface soils and waste materials in this area
is present as the mineral scorodite, ferric arsenate, which
is highly insoluble and thus of low bioavailability/
bioaccessibility.
Other sources of heavy metal contaminants in agri-
cultural soils have been well documented and include
phosphatic fertilizers (Cd), sewage sludge and pig slurry
(Cd, Pb and Cu) and a wide range of industrial and
urban emissions. Their sources, exposure pathways,
absorption, retention and toxic effects on grazing live-
stock are discussed by Underwood and Suttle (1999).
To implement the new ‘‘contaminated land regime’’
guideline values for metals in soils, derived as a result of
strategies based on risk assessment, will soon be pub-
lished as a means of deciding whether land may be
regarded as contaminated or not. It is as yet uncertain
as to how these values will affect the quality and value
of agricultural land and whether these will be applied
regarding the suitability of land for livestock farming.
One problem with deriving guidelines is that they are
usually based on total metal concentrations in the soil and
do not take into account different chemical and mineral
forms which naturally influence the metal’s solubility,
mobility and bioavailability. Similarly, they do not take
into account the fact that metals present as contaminants
in soils may change their form and availability over time
as a result of ageing and transformation processes.The majority of the livestock-related research sum-
marised in this article refers to work undertaken in the
UK and the developed world. Appleton (1992) has
reviewed the use of regional geochemical maps, some
compiled for mineral exploration, in developing countries
to identify areas of potential mineral deficiencies and
excesses in cattle. Attention is drawn to (a) relations
between areas of low Cu, Mn and Zn indicated by
regional geochemical maps for Swaziland (Forgeron,
1979) and possible deficiencies of these elements in
grazing livestock previously proposed as a result of
analysis of soils, herbage, cattle liver and serum in themiddle veld topographic zone (Ogwang, 1988); (b)
broad agreement between soil and drainage geochem-
istry (Stephenson et al., 1983) showing the potential for
deficiencies of Co and Cu and for Mo-induced Cu defi-
ciency in a mixed farming region in the centre of northern
Sumatra; (c) areas in which geochemical maps for NE
Zimbabwe (Dunkley, 1987) indicate the likelihood of
deficiencies of Co, Cu, Mn and Zn; (d) possible animal
deficiencies in extensive areas underlain by Precambrian
crystalline rocks of low Co, Cu, Mn and Zn shown by
the geochemical atlas of eastern Bolivia (Appleton and
Llanos, 1985) and (e) extensive areas of northern Sierra
Leone with low levels of Co, Cu and Mn in drainage
sediments (MacFarlane et al., 1981), again mainly
underlain by Precambrian crystalline rocks, in which
cattle are reared by nomadic farmers but where cattle
could be reared more intensively in the future.
A follow up report prepared by the Centre for Tropical
Veterinary Medicine, Edinburgh (1992), provided a
review of published literature on the mineral status of
livestock and pastures in these tropical countries. Infor-
mation was found to be sparse for the specific areas
covered by the regional geochemical maps reported by
Appleton (1992), and it was recommended that sampling
Table 1
Mean contribution of soil to total daily intake of As and Cu in 11 herds of cattle in Cornwall, England
Farm type
(based on soil As)
Mean % soil
ingested
Soil (mg kg1) Total daily intake (mg d1) % element ingested as soil
As Cu As Cu As Cu
20–50 mg kg1 As 1.4 24 26 7 140 67 10
n=3
55–140 mg kg1 As 1.4 85 93 21 175 76 10
n=3
160–259 mg kg1 As 1.1 202 199 52 190 58 16
n=5
I. Thornton / Applied Geochemistry 17 (2002) 1017–1028 1023
7/21/2019 Geochemistry and the Mineral Nutrition of Agricultural Livestock and Wildlife
http://slidepdf.com/reader/full/geochemistry-and-the-mineral-nutrition-of-agricultural-livestock-and-wildlife 8/12
of soil, forage, animal fluid and tissue should now be
carried out in selected areas to further assess the usefulness
of geochemical maps to identify areas where trace element
deficiencies or excesses might affect cattle productivity
under tropical conditions. On the basis of limited
correlations between reported deficiencies in livestock
and data provided by the geochemical maps, the authorsproposed levels of Cu, Co, Mn and Zn in drainage
sediments below which deficiencies in unsupplemented
grazing ruminants might occur.
More recent research in East Africa has demonstrated
the role of geochemistry in identifying trace element
imbalances in pasture in Zimbabwe and Kenya (Plant et
al., 1996; Fordyce et al., 1996; Jumba et al., 1996a,b).
However, in NE Zimbabwe, where stream sediment
geochemical maps provided useful indication of levels of
Zn and Cu in soil and grass, there was no correlation
with levels in bovine serum. The authors concluded that
this was probably due to differences in the species andmaturity of forages sampled and in the gender and age
of cattle from which serum had been obtained (Fordyce et
al., 1996). In Kenya, mineral concentrations in pastures
frequently fall below published standards for grazing
livestock. However, it is suggested that levels of Zn and
P below these standards do not necessarily imply that
production will benefit by supplementation; requirements
for livestock production may well be met. It was noted
that there were large differences in Zn concentrations
between herbage species sampled (Jumba et al., 1996b).
3. Geochemistry and wildlife nutrition
Further to the successful application of geochemistry
to agriculture and the recognition that geochemical
maps can delineate areas in which food crop and farm
livestock production and health could be influenced by
trace element deficiencies and excesses, the People’s
Trust for Endangered Species commissioned research to
see if this approach could be applied to wildlife con-
servation areas in Kenya. Here, a potential problem had
been recognised by the Kenyan Department of Wildlife
Conservation and Management within the Lake
Nakuru National Park. It was designated as the firstsanctuary for the black rhinoceros, an animal in danger
of extinction as a result of poaching. In this region in
the Rift Valley there is a history of mineral deficiencies
of ranched cattle and both the impala and waterbuck
within the Park show signs of possible deficiencies. The
impala show weakness and stiffness of the back legs and
the waterbuck, emaciation and loss of coat colour
(Maskall and Thornton, 1989).
The problem is that, in order to support conservation
and encourage tourism, wildlife are now enclosed in
what can be described as island parks. Instead of being
able to roam over very large areas, the animals are now
dependent on herbage and browse plants in relatively
small areas. Lake Nakuru National Park is only 160
km2. Some 20 black rhinoceros have been moved into
the Park, which is also the home of the relatively rare
Rothchilds giraffe.
A reconnaissance geochemical survey was carried out
to establish the mineral status of soils and selected plantspecies and the results related to the health of the animals
(Maskall and Thornton, 1989). Soil and plant samples
collected on a one-kilometre grid were analysed for 25
elements using ICP-AES and blood samples analysed
for Cu and vitamin B12.
The total concentrations of Cu and Co in soils were
low, a geochemical feature shared by many of the Rift
Valley soils derived from volcanic ash sediments and
other volcanic rocks. Total soil Se and P levels were also
relatively low. Grass species contained higher levels of
Cu and Co and lower levels of Se compared to browse
plants. Molybdenum levels in all plants reached rela-tively high values and the availability of this element
appeared to increase in wetter soils of high pH near the
lake shore (Maskall and Thornton, 1991, 1996). Over
30% of impala sampled had blood Cu levels below that
regarded adequate for domestic animals and were sig-
nificantly lower than those typical for impala throughout
Kenya. The relatively high Mo content of grasses and
browse plants is believed to contribute to possible Cu
deficiencies of impala and water buck in the park. The
lack of data on the mineral requirements of other wildlife
species prevents an assessment of the risk of deficiencies to
these species, including rhino, at the present time. As a
result of this work, recommendations were made to the
Kenya Department of Wildlife Conservation and Man-
agement that mineral salts containing Co, Cu and Se
should be made available to wildlife in Lake Nakuru
Park.
Further geochemical studies showed similar low con-
centrations of Cu and Co in brown clay loams at the Ol
Ari Nyiro Wildlife Reserve and the dark brown clays
developed from lavas from Mount Kenya at Solio
Wildlife Reserve. Soils derived from basaltic lavas at
Aberdares Salient, Amboseli National Park and basalts
at Lewa Downs Wildlife Reserve had higher concen-
trations of trace elements reflecting the basic nature of the parent material (Maskall and Thornton, 1991).
Black rhino at Solio had lower blood Cu levels than
rhino sampled in Zimbabwe and may indicate Cu defi-
ciency in the former. Jumba et al (1996a,b) reported low
‘‘total’’ concentrations of Ca, Mg, P, Cu and Zn in her-
bage from the Mount Elgar region of Kenya. Recent
geochemical studies in the Shimba Hills National Park,
home of the endangered Roosevelt sable, are reported
elsewhere in this volume.
Bowell and Ansah (1993) studied soils and vegetation
in the Mole National Park, Ghana and found Fe, Mn
and Co to be mainly fixed in the mineral fraction of the
1024 I. Thornton / Applied Geochemistry 17 (2002) 1017–1028
7/21/2019 Geochemistry and the Mineral Nutrition of Agricultural Livestock and Wildlife
http://slidepdf.com/reader/full/geochemistry-and-the-mineral-nutrition-of-agricultural-livestock-and-wildlife 9/12
soil, while Cu, Mo and Se were readily extractable. They
also showed Cu, Co and Mn to be present in higher
concentrations in grasses and Mo and Se in browse
plants.
The low Cu status of soils found in a geochemical
reconnaissance survey of the Mkomazi Game Reserve in
Tanzania suggest a possible Cu deficiency problem inherbivores (Abrahams and Bowell, 1998). McNaughton
(1990) proposed that seasonal movements of migrating
grazers in the Serengeti National Park in Tanzania were
influenced by the need for minerals, and in particular
Ca, Na, N, Cu and Zn, together with Mg and P for
lactating females and young animals. Of course, there
are numerous other situations when this type of
approach could be applied.
However, it has long been recognised that wildlife
regularly and intentionally eat soil from specific loca-
tions, commonly referred to as salt licks (Ayeni, 1971).
The reason for this geophagia has remained obscurethough it has commonly been supposed that this soil is
ingested to supplement minerals deficient in the animal’s
diet. Studies aimed at identifying a common nutritional
feature of these lick soils have proven inconclusive,
though soil enrichment with Na has been noted (Kreulen
and Jager, 1984; Abrahams, 1999) and it has been sug-
gested that this mineral is the main attraction (Tracy
and McNaughton, 1995).
Osteophagia or bone consumption has also been
noted in giraffe at the Timbavati Private Nature
Reserve, South Africa (Langman, 1978) and in sable at
Shimba Hills National Reserve, Kenya (Sekulic, 1977)
as a probable source of Ca and/or P.
4. The future
Thornton (1993) referred to the fact that geochemical
maps and atlases continue to cover greater areas and
would provide an exciting opportunity to study the global
significance of many of the problems previously studied
on a local or regional scale. The group of European
Geological Surveys (FOREGS, 1996) has made con-
siderable progress in bringing together existing geo-
chemical survey information Europe-wide and planshave been drawn up for further sampling strategies to
complete the European picture. On a global scale, an
updated proposal for a geochemical reference network
has been published, with the prime aim of providing a
framework of systematic baseline data to facilitate the
preparation of a World Geochemical Atlas. Both the
European mapping programme and the worldwide
initiative would have obvious applications to agri-
culture, including animal health and production (Darn-
ley et al., 1995).
There is also obvious potential on a regional and
national scale for the consolidation and interpretation
of previous geochemical survey information based on
stream sediment, soil and vegetation sampling—fre-
quently undertaken for purposes of mineral exploration.
One of the unfortunate limitations of applying these
data for environmental purposes is that frequently the
analytical detection limits employed for exploration
purposes are too high to be meaningful at the levelrequired for the identification of animal deficiencies.
It has already been noted that there have been
encouraging developments in the application of regional
geochemical surveys to livestock nutrition in developing
countries. It is emphasised that there is an urgent need for
greater data sources on veterinary/agricultural information
for comparison with these geochemical maps/atlases in the
tropics. The opportunities for multi-disciplinary research in
the developing world are enormous and call for the
establishment of teams encompassing geochemists, soil
scientists, biologists and veterinary scientists together
with those agencies responsible for agricultural andwildlife development.
The enormity of the problems ahead were noted by
Mills (1985), who concluded that ‘‘despite increasing
experimental evidence that anomalies in trace element sup-
ply can influence the growth, reproduction performance or
immunocompetence of domestic animals, few data exist
from which the incidence and economic significance of
such problems can reliably be assessed. In particular,
difficulties are experienced in assessed economic losses
arising from disorders giving rise to non-specific effects
on growth, fertility and resistance to infection induced
by deficiencies of Cu, Co, Zn or Se or by Mo excess’’. If
one considers Cu deficiency alone, in the 1980s it was
estimated ‘‘that characteristic clinical evidence of Cu
deficiency probably develop annually in about 0.9% of
the UK cattle population’’. Mills (1985) suggested that
‘‘assuming very conservatively, that similar conditions
govern the frequency of this disorder worldwide, the
approximate magnitude of the problem could be that
11.3106 clinically identifiable cases of severe Cu defi-
ciency develop in cattle annually. This estimate takes no
account of losses in buffalo, goats and other ruminants’’.
The extent of the problem is illustrated in Table 2 which
lists those tropical countries in which Cu deficiency in
cattle and sheep was recognised as a serious problem inthe 1980s (Shorrocks and Alloway, 1985). Undoubtedly,
more information is now available. Mills further suggests
‘‘that losses from severe Co deficiency may be similar in
magnitude’’. Mills specifically draws attention to the
problems in developing countries and to the problem of
focussing on inorganic element nutrition in relation to
animal productivity when a complex interplay of multiple
deficiencies and infections confuses the clinical picture.
The situation in central and eastern Europe warrants
special consideration as, until relatively recent political
change, there had been difficulties in obtaining reliable
information on agricultural problems and their extent and
I. Thornton / Applied Geochemistry 17 (2002) 1017–1028 1025
7/21/2019 Geochemistry and the Mineral Nutrition of Agricultural Livestock and Wildlife
http://slidepdf.com/reader/full/geochemistry-and-the-mineral-nutrition-of-agricultural-livestock-and-wildlife 10/12
severity. A notable exception was work on biogeochemical
mapping in the USSR by scientists of the Vernadsky Insti-
tute, Moscow, based on extensive soil sampling. This lead
to the recognition of zones and zonal provinces in which
groups of trace and major elements were present at low,
adequate or high concentrations and in which animals
showed a biological response (Kovalsky, 1970, 1979;
Kovalsky and Andreanova, 1970). This valuable
compilation of data was regrettably not followed up by
more detailed investigations or, if so, these have not
been made available to the wider scientific community.
There is obvious potential for building on this geo-
chemical baseline.
The growing problem of contaminated land and the
development of regulations at a national, European and
international level has jointly focussed on the protection
of human health and of ecosystem as noted earlier in
this paper. In particular, the impacts of historical and
more recent mining and smelting activities have left a
legacy of extensive areas of agricultural and natural
land contaminated with one or more heavy metals,
sometimes at very high concentrations. An example of the worldwide relevance of this is illustrated by recent
studies in Korea (Jung and Thornton, 1996). The issue
of contaminated land and its potential impacts is again
of paramount importance in central and eastern Europe.
The use of land for livestock production will no doubt
be influenced by further developments in this regulatory
process, in that much land heavily contaminated by
potentially toxic metals is used for livestock rearing at
this time and could, in certain cases, lead to the pro-
duction and sale of livestock products, particularly liver
and kidney, that exceed recommended levels for human
consumption.
The applications of geochemistry to wildlife nutrition
and conservation is still relatively new and the recent
work in Tanzania and Kenya is encouraging. There is,
however, a huge potential to develop this type of inves-
tigation further, particularly in southern Africa and
Latin America. Funding such work must surely be the
responsibility of international and national developmentagencies with a view to supporting the future growth of
tourism based on successful wildlife management.
Finally, it is recognised that the geochemist and chemist
now have more sophisticated tools to facilitate the
understanding of processes in the soil–plant–animal
system that influence and control the supply of essential
nutrient elements and/or exposure to potentially toxic
metals. Recent developments in sampling and analytical
procedures have focussed on the speciation of elements
in the soil and soil solution as the first critical stage in the
source–pathway–animal receptor model. It is envisaged
that these analytical tools, coupled with modellingtechniques and the use of GIS will lead to improved
methods of applying geochemical maps to livestock and
wildlife nutrition and sustainability.
In conclusion, it is stressed that geochemists must not
work in isolation. If full use is to be made of geochemical
data, whatever the scale, collaboration with agricultural
and veterinary scientists and with those responsible for
wildlife management is essential if meaningful progress
is to be made.
References
Abrahams, P.W., 1999. The chemistry and mineralogy of three
Savannah lick soils. J. Chem. Ecol. 25, 2215–2228.
Abrahams, P.W., Bowell, R.J., 1998. Soil geochemical mapping
of Mkomazi. In: Coe, M.J., McWilliam, N.C., Stone, G.N.,
Packer, M.J. (Eds.), Mkomazi: The Ecology, Biodiversity
and Conservation of a Tanzanian Savannah. Royal Geo-
graphical Society (with The Institute of British Geo-
graphers), London, pp. 25–39.
Abrahams, P.W., Thornton, I., 1987. Distribution and extent
of land contaminated by arsenic and associated metals in
mining regions of south-west England. Trans. Inst. Min.
Metal. (Sect. B: Appl. Earth Sci.) 96, B1–B8.Abrahams, P.W., Thornton, I., 1994. The contamination of
agricultural land in the metalliferous province of southwest
England: implications to livestock. Agric. Ecosys. Environ.
48, 25–137.
Appleton, J.D., 1992. Review of the Use of Regional Geo-
chemical Maps for Identifying Areas where Mineral Defi-
ciencies or Excesses May Affect Cattle Productivity in
Tropical Countries (Technical Report WC/92/24). British
Geological Survey, Keyworth, Nottingham.
Appleton, J.D., Llanos, A., 1985. Geochemical Atlas of East-
ern Bolivia. British Geological Society.
Ayeni, J.S.O., 1971. Mineral licks—a literature review. Obeche
7, 46–53.
Table 2
Tropical countries in which Cu deficiency in grazing cattle and
sheep is a serious problem
Argentina Malaysia
Bolivia Malawi
Brazil Mexico
Colombia Panama
Costa Rice Peru
Cuba Philippines
Dominican Rep. Senegal
Ecuador South Africa
El Salvador Sudan
Ethiopia Surinam
Guatemala Swaziland
Guyana Tanzania
Haiti Trinidad
Honduras Uruguay
India Venezuela
Indonesia Zaire
Kenya Zimbabwe
Saudi Arabia
Taken from Shorrocks and Alloway, 1985.
1026 I. Thornton / Applied Geochemistry 17 (2002) 1017–1028
7/21/2019 Geochemistry and the Mineral Nutrition of Agricultural Livestock and Wildlife
http://slidepdf.com/reader/full/geochemistry-and-the-mineral-nutrition-of-agricultural-livestock-and-wildlife 11/12
Bowell, R.J., Ansah, R.K., 1993. Trace element budget in an
African savannah ecosystem. Biogeochemistry 20, 103–126.
Brebner, J., Suttle, M.F., Thornton, I., 1987. Assessing the
availability of ingested soil cobalt for the synthesis of vitamin
B12 in the ovine rumen. Proc. Nutr. Soc. 46, 766A.
Centre for Tropical Veterinary Medicine (CTVM) Edinburgh Uni-
versity, 1992. Report on the Mineral Status of Animals in some
Tropical Countries and their Relationship to Drainage Geo-
chemical Maps of Minerals in those Countries (Technical Report
WC/92/60). British Geological Survey, Keyworth, Nottingham.
Colbourn, P., Thornton, I., 1978. Lead pollution in agricultural
soils. J. Soil Sci. 29, 513–526.
Darnley, A.G., Bjo ¨ rklund, A., Bølviken, B., Gustavsson, N.,
Koval, P.V., Plant, J.A., Steenfelt, A., Tauchid, M., Xuejing,
Xie, 1995. A Global geochemical database for environmental
and resource management. UNESCO Publishing, Paris.
Dunkley, P.N., 1987. A Regional Geochemical Exploration
Survey of the Makaha Area, North-east Zimbabwe (Open
File Rep. Br. Geol. Survey). Overseas Directorate, MP/87/17.
Fordyce, F.M., Masara, D., Appleton, J.D., 1996. Stream
sediment, soil and forage chemistry as indications of cattlemineral status in north east Zimbabwe. In: Appleton, J.D.,
Fuge, R., McCall, G.J.H. (Eds.), Environmental Geochem-
istry and Health. Geological Society Special Publication No.
113, pp. 23–37.
FOREGS, 1996. Report Geochemistry Task Group 1994–1996.
Forum of European Geological Surveys (Technical Report
WP/95/14). British Geological Survey, Keyworth, Nottingham.
Forgeron, D., 1979. Regional Stream Sediment Geochemical
Reconnaissance of Swaziland. Bull. 8.. Geological Survey
and Mines Department, Swaziland.
Geeson, M.A., Abrahams, P.W., Murphy, M.P., Thornton, I.,
1998. Fluorine and metal enrichment of soils and pasture
herbage in the old mining areas of Derbyshire, UK. Agric.
Ecosys. Environ. 68, 217–231.
Healy, W.B., 1968. Ingestion of soil by dairy cows. N.Z. J.
Agric. Res. 11, 487–499.
Jumba, I.O., Suttle, N.F., Wandiga, S.O., 1996a. Mineral
composition of tropical forages in the Mount Elgon region of
Kenya. 1. Macro-minerals. Trop. Agric. 73, 108–112.
Jumba, I.O., Suttle, N.F., Wandiga, S.O., 1996b. Mineral
composition of tropical forages in the Mount Elgon region of
Kenya. 2. Trace elements. Trop. Agric. 73, 113–118.
Jung, M.C., Thornton, I., 1996. Heavy metal contamination of
soils and plants in the vicinity of a lead-zinc mine, Korea.
Appl. Geochem. 11, 53–59.
Kovalsky, V.V., 1970. The geochemical ecology of organisms
under conditions of varying contents of trace elements in theenvironment. In: Mills, C.F., Livingstone, S. (Eds.), Trace
Element Metabolism in Animals. Edinburgh and London,
pp. 385–397.
Kovalsky, V.V., 1979. Geochemical ecology and health prob-
lems. Phil. Trans. R. Soc. Lond. B288, 185–191.
Kovalsky, V.V., Andreanova, G.A., 1970. Trace Elements in
Soils of the USSR. Hayka, Moscow. (in Russian).
Kreulen, D.A., Jager, T., 1984. The significance of soil inges-
tion in the utilization of arid rangelands by large herbivores
with special reference to natural licks on the Kalahari pans.
In: Gilchrist, F.M.C., Machie, R.I. (Eds.), Herbivore Nutri-
tion in the Subtropics and Tropics. The Science Press,
Johannesburg, pp. 204–221.
Langman, V.A., 1978. Giraffe pica behaviour and pathology as
indicators of nutritional stress. J. Wildlife Manag. 42, 141–
147.
Leech, A., Howarth, R.J., Thornton, I., Lewis, G., 1982. The
incidence of bovine copper deficiency in England and the
Welsh borders—the current situation. Vet. Rec. 111, 203–204.
Leech, A.F., Thornton, I., 1987. Trace elements in soils and
pasture herbage on farms with bovine hypocupraemia. J.
Agric. Sci. 108, 591–597.
MacFarlane, A., Crow, M.J., Arthura, J.W., Wilkinson, A.F.,
Aucott, J.W., 1981. The Geology and Mineral Resources of
Northern Sierra Leone (No. 7). British Geological Survey.
Maskall, J.E., Thornton, I., 1989. The mineral status of Lake
Nakuru National Park, Kenya: a reconnaissance survey. Afr.
J. Ecol. 27, 191–200.
Maskall, J.E., Thornton, I., 1991. Trace element geochemistry
of soils and plants in Kenyan conservation areas and impli-
cations for wildlife nutrition. Environ. Geochem. Health 13,
93–107.
Maskall, J.E., Thornton, I., 1996. The distribution of trace and
major elements in Kenyan soil profiles and implications forwildlife nutrition. In: Appleton, J.D., Fuge, R., McCall,
G.J.H. (Eds.), Environmental Geochemistry and Health.
Geological Society Special Publication 113, pp. 153–161.
McGrath, S.P., Loveland, P.J., 1992. The Soil Geochemical Atlas
of England and Wales. Blackie Academic and Professional.
McNaughton, S.J., 1990. Mineral nutrition and seasonal
movements of African migratory ungulates. Nature 345,
613–615.
Mills, C.F., 1979. Trace elements in animals. Phil. Trans. R.
Soc. Lond., B 288, 51–63.
Mills, C.F., 1985. Changing perspectives in studies of the trace
elements and animal health. In: Mills, C.F., Bremner, I.,
Chesters, J.K (Eds.), Proc. 5th Internat. Symp. Trace Elements
in Man and Animals (TEMA 5). Commonwealth Agricultural
Bureaux, Farnham Royal, Aberdeen, UK, pp. 1–9.
Oelschlager, W.K., Loeffler, K., Opletova, I., 1970. Retention
of fluoride in bones and bone sections of oxen in respect of
equal increases in fluorine in the form of soil, flue dust from
an aluminium reduction plant and sodium fluoride. Land-
wirsch. Forsch 23, 214–224.
Ogwang, B.H., 1988. The mineral status of soil, forage and
cattle tissues in the middleveld of Swaziland. Exper. Agric.
24, 177–182.
Patterson, J.B.E., 1938. Some observations on a disease of
sheep on Dartmoor. Emp. J. Expt. Agric. 6, 262.
Phillips, C.J.C., Foster, C., Morris, P., Teverson, R., 2000. The
Role of Cattle Husbandry in the Development of a Sustain-able Policy to Control M. bovis Infection in Cattle. Report to
the Ministry of Agriculture, Fisheries and Food. Veterinary
Laboratories Agency, pp. 30–31.
Plant, J.A., Baldock, J.W., Smith, B., 1996. The role of geo-
chemistry in environmental and epidemiological studies in
developing countries: a review. In: Appleton, J.D., Fuge, R.,
McCall, G.J.H. (Eds.), Environmental Geochemistry and
Health. Geological Society Special Publication No. 113, pp.
7–22.
Rieuwerts, J.S., Thornton, I., Farago, M.F., Ashmore, M.R.,
1998. Factors influencing metal bioavailability in soils: pre-
liminary investigations for the development of a critical
approach for metals. Chem. Spec. Bioavail. 10, 61–75.
I. Thornton / Applied Geochemistry 17 (2002) 1017–1028 1027
7/21/2019 Geochemistry and the Mineral Nutrition of Agricultural Livestock and Wildlife
http://slidepdf.com/reader/full/geochemistry-and-the-mineral-nutrition-of-agricultural-livestock-and-wildlife 12/12
Sekulic, R., 1977. Some aspects of behaviour, ecology and
conservation of the sable (Hippotragus niger (Harris, 1838)
and roan (Hippotragus equinus (Desmarest, 1804) antelopes
in the Shimba Hills National Reserve, Kenya. BA Hons dis-
sertation. Harvard University, USA.
Shorrocks, V.M., Alloway, B.J., 1985. Copper in Plants, Ani-
mals and Human Nutrition. Copper Development Associ-
ation, Potters Bar, UK.
Smart, L., 1991. The Agricultural Implications of Anomalous
Molybdenum Concentrations in Reclaimed Saltmarsh Soils
along the Suffolk Coast. Unpublished MSc thesis, Imperial
College of Science, Technology and Medicine, London.
Stephenson, B., Ghazali, S.A., Widjaja, H., 1983. Regional
Geochemical Atlas Series of Indonesia: 1. Northern Suma-
tra. British Geological Survey.
Suttle, N.F., Alloway, B.J., Thornton, I., 1975. An effect of soil
ingestion and utilisation of dietary copper by sheep. J. Agric.
Sci. 84, 249–254.
Suttle, N.F., Abrahams, P., Thornton, I., 1984. The role of
soildietary sulphur interaction in the impairment of copper
absorption by ingested soil in sheep. J. Agric. Sci. Cambridge103, 81–86.
Suttle, M.F., Brebner, J., Hall, J., 1991. Faecal excretion and
retention of heavy metals in sheep ingesting top soil from
fields treated with metal rich sludge. In: Mouncilovic, B.
(Ed.), Proc. 7th Internat. Symp. on Trace Elements in Man
and Animals (TEMA 7). Dubrovnik, IMI, Zagreb, pp. 32-7–
32-8.
Thomson, I., Thornton, I., Webb, J.S., 1972. Molybdenum in
black shales and the incidence of bovine hypocuprosis. J. Sci.
Food Agric. 23, 871–891.
Thornton, I., 1974. Biogeochemical and soil ingestion studies in
relation to the trace element nutrition of livestock. In:
Hoekstra, W.G., Suttie, J.W., Ganther, H.E., Mertz, W.
(Eds.), Trace Element Metabolism in Animals 2. University
Press, Baltimore, MD, pp. 451–454.
Thornton, I., 1983. Soil–plant–animal interactions in relation
to the incidence of trace element disorders in grazing live-
stock. In: Suttle, N.F., Gunn, R.G., Allen, W.M., Linklater,
K.A., Wiener, G. (Eds.), Trace Elements in Animal Produc-
tion and Veterinary Practice. British Society of Animal Pro-
duction, Edinburgh, pp. 39–49.
Thornton, I., 1993. Environmental geochemistry and health in
the 19900s: a global perspective. Appl. Geochem. 2 (Suppl.),
203–210.
Thornton, I., Abrahams, P.W., 1983. Soil ingestion—a major
pathway of heavy metals into livestock grazing contaminated
land. Sci. Tot. Environ. 28, 287–294.
Thornton, I., Kinniburgh, D., 1979. Intake of lead, copper and
zinc by cattle from soil and pasture. In: Kirchgessner, M.
(Ed.), Trace Element Metabolism in Man and Animals, vol.
3. Technische Universita ¨ t Mu ¨ nchen, Freising, Weihen-
stephan, p. 499.
Thornton, I., Webb, J.S., 1970. Geochemical reconnaissance
and the detection of trace element disorders in animals. In:
Mills, C.F. (Ed.), Trace Element Metabolism in Animals.
Livingstone, Edinburgh and London, pp. 397–407.
Thornton, I., Webb, J.S., 1979. Geochemistry and health in the
United Kingdom. Phil. Trans. R. Soc., London 288, 151–
168.Tracy, B.F., McNaughton, S.J., 1995. Elemental analysis of
mineral lick soils from the Serengeti National Park, the
Konza Prairie and Yellowstone National Park. Ecography
18, 91–94.
Underwood, E.J., 1962. Cobalt. In: Trace Elements in Human
and Animal Nutrition, second ed. Academic Press, New
York and London, pp. 123–156.
Underwood, E.J., 1966. The Mineral Nutrition of Livestock.
Commonwealth Agricultural Bureau.
Underwood, E.J., Suttle, N.F., 1999. The Mineral Nutrition of
Livestock. Essentially toxic elements (Aluminium, Arsenic,
Cadmium, Fluorine, Lead, Mercury). CABI Publishing,
Wallingford. (Chapter 18).
Webb, J.S., 1964. Geochemistry and life. New Scient. 23, 504–
507.
Webb, J.S., Thornton, I., Thompson, M., Howarth, R.J.,
Lowenstein, P.L., 1978. The Wolfson Geochemical Atlas of
England and Wales. Oxford University Press.
1028 I. Thornton / Applied Geochemistry 17 (2002) 1017–1028