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The evolution of herbal medicine: Behavioural perspectives

Article in Animal Behaviour · November 2005

DOI: 10.1016/j.anbehav.2005.03.005

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 ANIMAL BEHAVIOUR, 2005, 70, 975–989
doi:10.1016/j.anbehav.2005.03.005

REVIEW
The evolution of herbal medicine: behavioural perspectives

BEN JA MI N L. H ART
School of Veterinary Medicine, University of California, Davis

(Received 12 December 2003; initial acceptance 13 February 2004;

final acceptance 7 March 2005; published online 3 October 2005; MS. number: ARV-27R)

The current popularity of traditional herbal supplements, coupled with recent findings that add scientific
legitimacy to the use of some medicinal herbs, prompts a question about the origins of herbal medicine
in animals and ancestral humans. Medicinal herbs are used by animals and humans with the apparent pro-
phylactic effects of reducing the likelihood or severity of illness from pathogens or parasites in the future.
Medicinal herbs with anti-inflammatory, antimicrobial, immunomodulatory and/or analgesic properties
are used in a therapeutic way to treat acute infections and inflammatory conditions, particularly in humans,
and could have proven lifesaving to individuals living in nature. Was the origin of such types of herbal med-
icine the result of animals and humans learning that specific plant parts are effective for preventing or treat-
ing certain maladies, or was the origin a result of natural selection for a behavioural predisposition to seek
out and use plant parts with particular physical or chemosensory markers of efficacy? Examining the pre-
dictions and requirements of both the learned and evolutionary explanations points primarily to an evolu-
tionary model for the origin of herbal medicine that was expanded and enhanced by learning and social
transmission. The evolutionary explanation accounts for the continued use of ineffective, as well as effec-
tive, medicinal herbs and the use of medicinal herbs with toxic effects. In animals one can point to origins
of the practice of herbal medicine, as well as other behavioural defences against pathogens and parasites, as
analogues of many aspects of modern human medicine and health care.
! 2005 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Although herbal medicinal products represent a fast
growth area of health-care products for both humans
and companion animals (Eisenberg et al. 1998; Bent &
Avins 1999), their use in humans appears to predate re-
corded history (Tyler 2000). Reviews of the earliest indica-
tions of herbal medicine often refer to the grave of
a Neanderthal man, dated 60 000 years ago, in which
were found pollen and flower fragments of different me-
dicinal plants (Solecki 1971). More recently, investigation
of the possessions found on the 5300-year-old ‘iceman’
discovered in 1991 in the Italian Alps reported pieces of
birch fungus, presumably used as a laxative and antibiotic
(Capasso 1998). Most herbal medicines of current interest
have come from ancient civilizations of Africa, the Asian
subcontinent, and North, Central and South America
(Phillipson 2001). Although use of medicinal herbs ap-
pears to be much more prevalent among humans than an-
imals, understanding the prominence in human use is
inextricably bound to perspectives gleaned from studies
of self-medication in animals and differences and similar-
ities between humans and animals in the types of herbal
medicine practised.
In this paper I will examine hypotheses to explain the
origins of herbal medicine in light of issues such as the
long delay between ingestion of a herb and the onset of
medicinal effects, the persistent use of medicinal herbs
that appear to be ineffective or toxic and the placebo
effect that undoubtedly permeates the practice of herbal
medicine as it does conventional medicine. I will explore
whether learning of some type, or natural selection, most
adequately explains the origin and maintenance of the
practice of the many forms of herbal medicine among
animals and humans.
BACKGROUND

It has been suggested that the practice of herbal medicine

can generally be divided into prophylactic and therapeutic
Correspondence: B. Hart, University of California, Department of Anat- uses (Phillips-Conroy 1986; Lozano 1998; Johns 1999).
omy, Physiology, and Cell Biology, School of Veterinary Medicine, One Given that illnesses vary from nondetectable to severe,
Shields Avenue, Davis, CA 95616, U.S.A. (email: blhart@ucdavis.edu). the use of medicinal herbs with regard to being
975
0003–3472/05/$30.00/0 ! 2005 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.
976 ANIMAL BEHAVIOUR, 70, 5
prophylactic or therapeutic lies along a continuum be-
tween these two types of uses. Thus, the designation of
a type of herbal medicine as prophylactic or therapeutic
may occasionally involve a grey area between the two types.
I will use the term ‘prophylactic’ to refer to the prevention
or reduction of the likelihood of illnesses or maladies in the
future in individuals not showing signs of illness and ‘ther-
apeutic’ to refer to use by individuals with an evident mal-
ady, such as an acute illness or flare-up of an inflammatory
process. The prophylactic use of herbs may be ongoing or
periodic. Periodic use would be expected, for example, if
the risk of illness or disease varies with season (Lozano
1998), whereas the therapeutic use would generally be
short term and restricted to the course of the illness.
In most instances, the reported use of medicinal herbs
by animals appears to be prophylactic (see Table 1). The
use of medicinal herbs in traditional human societies, on
the other hand, appears to be primarily for treatment of
active or acute conditions. At least, this is the sense con-
veyed in the literature on herbal medicines in traditional
societies (Perry & Metzger 1980; Iwu 1993; Ghazanfar
1994; van Wyk et al. 1997; Ross 1999, 2001; Blumenthal
et al. 2000; Chevallier 2000; Parrotta 2001; Williamson
2003; Khare 2004). A perusal of the indications in tradi-
tional societies for 25 medicinal herbs used by humans,
for example, reveals a frequent mention of acute infec-
tions or inflammatory conditions including abdominal
pain, vomiting, gastrointestinal infections, diarrhea, ulcer-
ative colitis, gastritis, intestinal worms, liver problems,
nausea, sore throat, arthritis, canker sores, toothache
and fever (Chevallier 2000; see Table 2).
For therapeutic treatment of a variety of acute infections
and inflammatory conditions in animals and ancient
humans, I propose that the most logical plant parts would
be those with constituents that are anti-inflammatory,
analgesic, antimicrobial and/or immunomodulatory.
These are properties that should be most effective in
reducing the severity or disabling effects of a variety of
infectious or inflammatory conditions.
The Question of Efficacy
Central to understanding the origin of either prophylactic
or therapeutic herbal medicine is the necessity for actual
efficacy of medicinal herbs. Although the hundreds of herbal
products recommended for long and varied lists of maladies
invite scepticism about efficacy, research over the past
couple of decades or so, conducted under current scientific
standards, provides evidence of the medicinal efficacy for
some herbal products especially for therapeutic effects.
Efficacy is most difficult to demonstrate for prophylactic
use because the behaviour is based on reducing the severity
or likelihood of adverse events in the future. Most evidence
of efficacy is circumstantial, involving correlation between
the occasions of use of herbs and times of high risk of the
disease under consideration, and laboratory tests of plant
parts for potency in controlling causative agents of the
disease under consideration. Table 1, which outlines ani-
mal examples of herbal medicine from data-based studies,
particularly illustrates prophylactic use. The consumption
of soil, dirt or clay (geophagy), seen in several herbivorous
and omnivorous mammals, is a type of self-medication
(Knezevich 1998), but is not included in Table 1 because
this type of self-medication does not involve plant parts.
Fur rubbing with millipedes by wedge-capped capuchin
monkeys, Cebus olivaceus (Valderrama et al. 2000) is another
type of self-medication that does not involve a plant prod-
uct and is similarly not included.
Demonstration of efficacy for the treatment of acute
infections is straightforward because medicinal herbs, or
extracts thereof, can be tested on laboratory animals with
induced conditions or on in vitro preparations to docu-
ment efficacy. In fact, tests on laboratory animals, and in
vitro preparations, are an essential part of the develop-
ment of modern medicines for federal approval. What is
largely missing for full demonstration of efficacy for the
treatment mode are multiple clinical trials with appropri-
ate control groups. As described below, Table 2 summa-
rizes evidence from laboratory animal and in vitro
studies of the efficacy of medicinal herbs used in the treat-
ment of disease conditions where the anti-inflammatory,
antimicrobial, immunomodulatory and/or analgesic prop-
erties were examined.
Prophylactic Use of Medicinal Herbs
The most substantial body of published research on
animal use of medicinal herbs is on whole-leaf swallowing
by chimpanzees, Pan troglodytes. Observations of wild
chimpanzees at several study sites reveal a behaviour in
which the animals periodically swallow whole leaves,
which pass through the intestinal tract intact. The behav-
iour is most frequently seen during the rainy season when
the risk of gastrointestinal nematode infection is highest
(Wrangham 1995; Huffman et al. 1996). In most instances,
the animals performing the behaviour reportedly show no
evident signs of acute illness. However, in one study, chim-
panzees seen swallowing whole leaves frequently had signs
of diarrhea and an increased tendency to rest or sleep
(Huffman et al. 1996). Whole leaves were found in faeces
along with expelled intestinal nematodes. It has been sug-
gested that parasites are purged by enhanced gastrointesti-
nal motility induced by whole-leaf swallowing (Huffman &
Caton 2001). Although the diarrhea and tendency to rest
and sleep suggest a therapeutic mode of herbal medicine,
the increased frequency of whole-leaf swallowing, during
the rainy season, suggests a prophylactic mode.
Another documented type of prophylactic use of me-
dicinal herbs is seen in nest fumigation by dusky-footed
wood rats, Neotoma fuscipes. Wood rats use the same stick-
houses over many seasons and even generations, so there
is a risk of build-up of nestborne ectoparasites, especially
fleas. Wood rats bring into their stickhouses leafy sprigs
of California bay, Umbellularia californica, and toyon, Het-
eromeles arbutifolia, along with sprigs of a long-recognized
food staple, live oak, Quercus spp. Bay leaves are found sig-
nificantly more often near the sleeping nest than are other
plants. In in vitro experiments, bay leaves reduced survival
of flea larvae to about 25%, compared to 80–90% survival
for larvae exposed to leaves of oak, toyon or no leaves
 REVIEW 977

Table 1. Examples of therapeutic and prophylactic use of medicinal herbs in animals

Animal species Plant species Marker Postulated effect Evidence of effect References

Bitter pith chewing

Chimpanzees, Vernonia sp. Bitter taste Treat illness (therapeutic) Correlation with Huffman & Seifu 1989
Pan troglodytes illness
Antimicrobial Huffman et al. 1993
extracts
Whole-leaf swallowing
Chimpanzees Aspilia sp. Rough surface Expulsion of tapeworms Correlation with risk Wrangham 1995
(prophylactic/therapeutic)
Chimpanzees Rubia sp. Bristly surface Expulsion of nematodes Increased gut Huffman et al. 1996;
(prophylactic/therapeutic) mortality Huffman & Caton 2001
Bonobos, Pan Manniophyton sp. Expulsion of nematodes Correlation with risk Dupain et al. 2002
paniscus (prophylactic/therapeutic)
Nest fumigation
Starlings, Agrimonia sp., Volatiles Killing nestborne mites, In vitro tests Clark & Mason 1985
Sturnus vulgaris Daucus sp. lice (prophylactic)
Wood rats, Umbellurlaria sp. Volatiles Killing nestborne fleas, In vitro tests Hemmes et al. 2002
Neotoma fuscipes (prophylactic)
Fur rubbing
Capuchin Unspecified leaves Volatiles Repel ectoparasites; control Correlation with risk Baker 1996
monkeys, bacteria, fungi (prophylactic)
Cebus capucinus
White-nosed coatis, Trattinnickia sp. Volatiles Repelling ectoparasites Correlation with risk Gompper & Hoylman
Nasua naria (propylactic) 1993
Consumption of medicinal herbs
Baboons, Balanites sp. Control schistosomiasis Correlation with risk Phillips-Conroy 1986
Papio hamadryas (prophylactic)

(Hemmes et al. 2002). A recent study revealed that bay, as
well as toyon, has flea-repelling properties (A. Alvarado,
R. Hemmes & B. L. Hart, unpublished data).
Related uses of plants for nest protection have been
studied in European starlings, Sturnus vulgaris, which
weave foliage from herbal plants into the nest matrix
when eggs are being incubated. In laboratory tests, plants
selected by starlings suppressed the hatching of louse eggs
and arrested the development of nestborne mites (Clark &
Mason 1985, 1988).
Perhaps the most widespread prophylactic use of me-
dicinal herbs in humans is the use of spices in food
preparation. Evidence for the antimicrobial effects of
spices in reducing the growth of ingested foodborne
pathogens and in the preservation of meat, has been
well documented (Billing & Sherman 1998). A scenario
has been postulated in which ancient humans, living in
hot climates, who happened to add spicy plant parts to
meat dishes, suffered less from foodborne illness than
those who did not and perhaps were able to store food
longer. The spice-using people were then possibly health-
ier and were able to rear more healthy offspring.
A review of common names of some medicinal herbs (see
Table 2) below reveals some overlap between plant parts
that are used as spices and medicinal herbs with antimicro-
bial effects. Not surprisingly, traditional herbal gardens
maintained in the Middle Ages, and in modern times, typi-
cally include plants that serve as spices and medicines. The
use of spices in everyday food preparation could be consid-
ered a type of prophylactic human herbal medicine, and the
use of the same plant parts in larger amounts at one time
could be considered a therapeutic use of medicinal herbs.
Therapeutic Use of Medicinal Herbs
In contrast to the widespread therapeutic use of medic-
inal herbs in humans, the therapeutic use by animals
appears to be more limited. Perhaps the most convincing
animal example comes from observations on chimpanzees
that appeared ill. The animals were subsequently observed
ingesting the pith of young shoots of Vernonia amygdalina;
they meticulously removed the outer bark and leaves of
stems to chew and suck upon the extremely bitter pith
(Huffman & Seifu 1989; Huffman et al. 1993). Sesquiter-
pene lactones, with possible effects on intestinal hel-
minths, amoeba and bacterial pathogens, have been
isolated from V. amygdalina (Ohigashi et al. 1991; Jisaka
et al. 1992).
The lack of comparable observations of self-medication
by ill individuals of other nonhuman species, or even other
chimpanzees, raises the question about possible reasons for
the apparent infrequency with which such self-medication
for therapeutic purposes occurs in animals in nature. I can
think of three nonmutually exclusive possibilities. (1)
There is a low probability that an observer will be present
to observe and record the behaviour of an animal that is ill
seek out and consume medicinal herbs. (2) Social trans-
mission of the behaviour among individuals in a group
may be less likely in nonverbal animals than in humans
(see below). (3) Animals may actually become acutely ill less
frequently than humans, and consequently, only infre-
quently use medicinal herbs for therapeutic properties.
That nonhuman animals use medicinal herbs less for
therapeutic purposes is suggested by the perspective that
the evolution of the genus Homo involved a dietary shift to
 978
Table 2. Characteristics of 25 medicinal herbs tested in controlled laboratory tests

ANIMAL BEHAVIOUR, 70, 5

Taste in Study
Scientific name Common name Parts used nature* Documented medicinal effectsy typez References

Alpinia galanga Ginger, siamese Rhizomes, roots, Spicy Antimicrobial, immunomodulatory IV Haraguchi et al. 1996
ginger seeds
LA, IV Bendjeddou et al. 2003
LA Matsuda et al. 2003
Barberis vulgaris Barberry Root bark, berries Bitter Anti-inflammatory, other LA Ivanovska & Philipov
(antihistaminic) 1996
IV Shamsa et al. 1999
Boswellia serrata Frankincense Gum resin, bark Astringent Anti-inflammatory LA Safayhi et al. 1991
LA Krieglstein et al. 2001
Bridelia ferruginea Bududi, kisni fruit Stem bark, Astringent Anti-inflammatory, antimicrobial, LA Olajide et al. 1999
leaves, fruit analgesic IV Akinpelu & Olorunmola
2000
LA Olajide et al. 2000
IV Talla et al. 2002
LA Olajide et al. 2003
Camellia sinensis Tea Roots, leaves, Astringent Anti-inflammatory, immunomodulatory, LA Sur et al. 2001
buds other (antidiarrheal) IV Zvetkova et al. 2001
LA Besra et al. 2003
Cassia alata Candle bush, Leaves, stem bark, NA Anti-inflammatory, antimicrobial, LA Ibrahim & Osman 1995
ringworm bush root bark, flowers analgesic IV Khan et al. 2001
LA Villasenor et al. 2002
IV Somchit et al. 2003
Curcuma xanthorrhiza Javanese turmeric Rhizomes Spicy Anti-inflammatory, antimicrobial LA Claeson et al. 1993
IV Hwang et al. 2000
Echinacea species Echinacea, Leaves, roots Bitter Anti-inflammatory, LA Rehman et al. 1999
(augustifolia, purpura) coneflower immunomodulatory LA Mattace Raso et al. 2002
Entada abyssinica Abyssinia entada Roots, leaves, NA Anti-inflammatory, antimicrobial LA Fabry et al. 1996
stem bark IV Fabry et al. 1998
IV Freiburghaus et al. 1998
LA Olajide & Alada 2001
Evodia rutaecarpa Evodia Fruits Bitter Anti-inflammatory, analgesic LA Matsuda et al. 1998
LA Moon et al. 1999
LA Kobayashi 2003
Glycyrrhiza glabra Licorice Roots Sweet Antimicrobial, immunomodulatory LA Utsunomiya et al. 1995
LA Utsunomiya et al. 2000
Harpagophytum Devil’s claw Roots Bitter Anti-inflammatory, analgesic LA Lanhers et al. 1992
pocumbens IV Loew et al. 2001
IV Jang et al. 2003
LA Andersen et al. 2004
Hydrastis canadensis Goldenseal Rhizomes Bitter Antimicrobial, immunomodulatory IV Rehman et al. 1999
LA Scazzocchio et al. 2001
IV Hwang et al. 2003
IV Mahady et al. 2003
Hypericum perforatum St John’s Wort Flowers Bitter Anti-inflammatory, antimicrobial LA Schempp et al. 1999
IV Mattace Raso et al. 2002
IV Hubner 2003
 Nigella sativa Black seed, Seeds Spicy Anti-inflammatory, antimicrobial, LA, IV Haq et al. 1999
black cumin immunomodulatory, analgesic LA Abdel-Fattah et al. 2000
LA Salem & Hossain 2000
LA Al-Ghamdi 2001
IV Khan et al. 2003
Panax ginseng Ginseng Roots Sweet Antimicrobial, IV Lim et al. 2002
immunomodulatory IV Shin et al. 2002
Phyllanthus sp. Phyllanthus, Leaves, roots Bitter Anti-inflammatory, LA, IV Suresh & Vasudevan
(amarus, emblica) shatter stone immunomodulatory, analgesic 1994
LA Ihantola-Vormisto et al.
1997
LA Santos et al. 2000
LA, IV Kiemer et al. 2003
LA Raphael & Kuttan 2003
Punica granatum Pomegranate Fruit rind, seeds, Astringent Antimicrobial, immunomodulatory, LA Kurokawa et al. 1993
root bark other (antidiarrheal) LA Das et al. 1999
LA Gracious Ross et al. 2001
Scrophularia frutescens Figwort Leaves, stems Bitter Anti-inflammatory, antimicrobial LA Fernandez et al. 1996
IV Fernandez et al. 1998
Syzygium cumini Jambal Stem bark, leaves Astringent Anti-inflammatory, antimicrobial LA Muruganandan et al.
2001
IV Shafi et al. 2002
Tanacetum parthenium Feverfew Leaves, seeds, Bitter Anti-inflammatory, antimicrobial, LA, IV Jain & Kulkarni 1999
flowers analgesic IV, LA Williams et al. 1999
Urtica dioica Stinging nettle Leaves, rhizomes Astringent Anti-inflammatory, IV Galelli & Truffa-Bachi
immunomodulatory 1993
LA, IV Riehemann et al. 1999
Viscum album European mistletoe Leaves, stems NA Antimicrobial, immunomodulatory IV Zarkovic et al. 2001
LA, IV Hajto et al. 2003
IV Karagöz et al. 2003
Withania somniferum Winter cherry, Roots, leaves Bitter Anti-inflammatory, immunomodulatory LA Al-Hindawi et al. 1992
Ashwagandha LA Ziauddin et al. 1996
LA, IV Davis & Kuttan 2000
Zingiber officinale Ginger Rhizomes Pungent Anti-inflammatory, antimicrobial LA, IV Agarwal et al. 2001
IV Thomson et al. 2002
LA Penna et al. 2003

*Taste in nature of the unprocessed plant part used was obtained from the references in the last column or one or more books on herbal medicines. In a few instances references to the taste
were not available (NA).
yEffects were reported in peer-reviewed, international journals focusing on the medicinal herb in question. In some instances where anti-inflammatory, antimicrobial, immunomodulatory or
analgesic effects were documented, other effects, if documented in references, were also mentioned. The sources of information on medicinal effects are listed under references.
zStudy types refer to analyses in laboratory animals (LA) or in vitro tests (IV).

REVIEW
979
980 ANIMAL BEHAVIOUR, 70, 5
increasing amounts of animal meat and fat (reviewed by
Johns 1999; Finch & Stanford 2004). This dietary shift,
corresponding to the evolution of a large brain (Kaplan &
Robson 2002), had the potential to result in a number of
health-eroding effects, including an increase in free radicals
from fat metabolism with a simultaneous reduction in in-
take of free-radical-scavenging antioxidants in natural veg-
etation (Johns 1990, 1999). The increased likelihood of
acute infections and inflammatory conditions in early hu-
mans could have set the stage for the natural selection of
the use of medicinal herbs high in antimicrobial, immuno-
modulatory and anti-inflammatory components.
Medicinal Herbs that are Anti-inflammatory,
Antimicrobial, Immunomodulatory
and/or Analgesic
It is reasonable to assume that a major emphasis of
herbal medicine use in ancestral humans was on treat-
ment of ailments such as inflammatory conditions or
acute infections that would have been threatening to an
individual’s immediate survival or welfare. The most
useful herbs would have been those whose unprocessed
parts would have anti-inflammatory, analgesic, immuno-
modulatory and/or antimicrobial effects.
Table 2 lists 25 medicinal herbs, each of which was the
focus of at least two data-based, laboratory studies con-
ducted since 1990 that provided evidence of efficacy
from anti-inflammatory, analgesic, antimicrobial and/or
immunomodulatory properties when the herbs were con-
sumed. Plants with antimicrobial and immunomodulatory
effects might expedite recovery from acute illness caused
by pathogens, and plants with anti-inflammatory and an-
algesic effects might alleviate the disabling aspects of in-
flammation in one or more organ systems, allowing the
individual to return to normal behaviour, including care
of dependent young. The studies considered as providing
evidence were experiments using laboratory rodents or in
vitro analyses.
Medicinal herbs considered as candidates for inclusion
in Table 2 were derived from literature searches using ‘me-
dicinal plants’ as a common term and searching under
each of the following as a second term: (1) anti-inflamma-
tory, (2) analgesic, (3) immunomodulatory, and (4) anti-
microbial. I next conducted a literature search on those
medicinal plants that met the subject criteria, using only
papers published in peer-reviewed, mainstream interna-
tional biomedical journals, including the four peer-re-
viewed journals devoted to medicinal herbs (Journal of
Ethnopharmacology, Phytomedicine Research, Planta Medica,
Fitoterapia). For each plant, I determined whether at least
two scientific, data-based, controlled experimental stud-
ies, with statistically significant results, had been pub-
lished in or after 1990 addressing one or more of the
types of effects named above. I also sought information
on the part of the plant used and its taste as found in na-
ture. Finally, I consulted five books authored by recog-
nized authorities for the plants’ taste in nature and parts
used (Ross 1999, 2001; Blumenthal et al. 2000; Chevallier
2000; Williamson 2003). Table 2 includes the scientific
and common name of the medicinal plants, part(s) typi-
cally used, reported taste of the unprocessed part (if this
information was available), type of documented medicinal
effect and references to the studies documenting efficacy.
Table 2 is far from exhaustive with regard to the medic-
inal plants and the scientific research papers that could
have been included. The list of references for each herb
was restricted to no more than five. References to research
reports of other medicinal herbs are available (Ross 1999,
2001; Blumenthal et al. 2000; Williamson 2003). I inten-
tionally excluded medicinal herbs used for behavioural ef-
fects such as alleviation of depression, enhancement of
cognitive function or behavioural arousal. In one in-
stance, however, I included a herb known primarily as
an antidepressant (St John’s Wort, Hypericum perforatum)
because there are reports documenting its anti-inflamma-
tory and antimicrobial effects, which could have predated
its use as an antidepressant.
Five of the medicinal herbs featured in Table 2 have been
subjected to systematic reviews or meta-analyses of multiple
clinical trials on humans using randomized, placebo-con-
trolled, double-blind, methodologies that met minimal cri-
teria for methodological quality. The effects of interest were
not necessarily the same effects reported in Table 2. These
medicinal herbs are Echinacea spp. (echinacea) for preven-
tion or treatment of the common cold (Melchart et al.
1995), H. perforatum (St John’s Wort) for treatment of mild
depression (Linde et al. 1996), Panex ginseng (ginseng) for
enhancement of physical and mental performance (Vogler
et al. 1999), Tancetum parthenium (feverfew) for prevention
of migraine (Ernst & Pittler 2000a) and Zingiber officiniale
(ginger) for treatment of nausea and vomiting (Ernst & Pit-
tler 2000b). The systematic reviews or meta-analyses re-
vealed inconsistent results with regard to efficacy.
Typically, some trials reported significant effects and some
nonsignificant effects, leading the authors of the reviews
or meta-analyses to conclude that at least some of the herbal
preparations in use are more effective than placebo.
Given the expected variability in potency of the herbal
plant products in nature, and variability in clinical con-
ditions of patients enrolled in such studies, inconsisten-
cies in reports of the efficacy of medicinal herbs might be
expected. That is, even if multiple clinical trials of sound
methodology had been conducted with each of the entries
of Table 2, focusing on the medicinal effects profiled in
the table, the results would probably be similar.
Table 2 does not include any putative medicinal herbs
for which ineffectiveness was documented. Undoubtedly
many putative medicinal herbs are no better than placebo,
but no purpose would be served by including such nega-
tive findings, even if they were available. I will deal with
implications of the variability in the potency of medicinal
plant products, including those that are ineffective or
toxic, under discussion of the implications of the evolu-
tionary explanation of the origin of herbal medicine.
Disclaimer
Although the research studies cited in Table 2 provide
evidence of efficacy for each of the medicinal plants listed,

this is not an endorsement of their use. Some medicinal
herbs currently marketed are available in concentrated
forms and could be toxic or have adverse interactions
with other medications (Ernst 1998). Even a medicinal
herb that is toxic may have been beneficial for ancient hu-
mans or may be beneficial for free-living animals con-
fronted with an acute illness. Such trade-offs can
generally be avoided with modern pharmaceutical
preparations.
ALTERNATIVE HYPOTHESES ON THE ORIGIN
OF HERBAL MEDICINE
In general, the two possibilities for the origins of the
prophylactic and the therapeutic types of herbal medicine
are learning and natural selection. Both explanations
assume that medicinal herbs are effective in preventing
or treating conditions at least some of the time. The
learning explanation seems to be favoured by most
authors dealing with the occurrence of herbal medicine
in animals (e.g. Sapolsky 1994; Huffman 1997, 2001; Loz-
ano 1998). Although not explicitly stated, the impression
conveyed by these authors is that individuals learn to
choose from a large array of plants, those plants or plant
parts that will be the most effective for a particular mal-
ady. Thus, the learning explanation applied to the pro-
phylactic use of herbal substances would specify that
individuals learn the association between the use of a spe-
cific medicinal herb (or group of herbs) and the future pre-
vention of pain or discomfort from a particular malady.
The learning explanation applied to the therapeutic use
would imply that if the use of a specific medicinal herb
(or group of herbs) was followed by alleviation of pain
and discomfort, the individual would use the same prod-
uct when that malady occurred again. The reduction of
pain or discomfort could result from resolution of the ill-
ness or from relief from inflammation.
The explanation based on natural selection for the
prophylactic use would specify an evolved predisposition
for individuals to seek out and use or consume plant
products that characterize one or more physical or chemo-
sensory markers, which in turn are correlated with
effectiveness in preventing or reducing a particular illness
in the future. As applied to the therapeutic use, this
explanation would specify an evolved predisposition for
sick individuals to seek out and consume plant products
with one or more chemosensory markers, and this would
lead to an increased likelihood of survival or earlier return
to normal activity. An increase in reproductive fitness is
assumed to be associated with the evolved predispositions.
The use of plant products for prevention or alleviation
of illness is a logical area in which to expect the behaviour
to be transmitted by social means, especially among
family or clan members. For social transmission to occur,
some individuals must first acquire the behaviour to serve
as ‘demonstrators’. Although social transmission of ele-
ments of herbal medicine would be important in the
dissemination of this behaviour, the behaviour must be
anchored to an initial learned or evolved acquisition.
Below I contrast different learning models with the
REVIEW 981
natural selection model to evaluate adequate explanations
of the origin of herbal medicine. After dealing with these
approaches, I then discuss social transmission.
The type of herbal medicine considered here is the use
of raw or nonprocessed plant parts that would be
available to animals or early humans in the natural
environment. A choice between the learning and evolu-
tionary explanations stems from the different predictions
and/or requirements of each. For the sake of discussion, I
contrast the learning and evolutionary explanations as
a dichotomy while acknowledging that, if an evolutionary
origin seems likely, learning undoubtedly interacted with
and reinforced the natural selection process. The relative
importance of natural selection and learning would be
reflected in the particular type of condition being affected
by herbal use.
Learning as a Basis for Herbal Medicine
One model of learning for the acquisition of herbal
medicine is a type of operant conditioning acquired
through appropriate reinforcement. Some contingency is
required between the performance of a behaviour, in this
case seeking out and using a specific plant or plant part,
and its reinforcement. For example, native Americans are
reported to have used bay leaves to repel mosquitoes (Balls
1962). One can imagine that the initial application of bay
leaves to the skin was accidental, but the distinctive aroma
would have remained on the skin and could have become
associated with the absence of mosquitoes. As another ex-
ample, many plant products have reinforcing excitatory,
calming or hallucinogenic effects that occur soon after in-
gestion and their use could be explained by learning.
For a behaviour such as seeking out and using a plant
part for medicinal effects to be learned using the conven-
tional operant paradigm, reinforcement must follow the
performance of the behaviour within a fairly short time;
this is usually expressed in seconds or minutes rather than
hours, days or weeks (Schwartz & Robbins 1995). Few
learned operants occur with one trial, so another general
requirement of operant conditioning is multiple sessions
of reinforced responses. Reinforcement during these mul-
tiple trials should also be more continuous than intermit-
tent; once learned, the behaviour can be maintained by
intermittent reinforcement.
Application of the operant learning model to the use of
herbs for the prevention or treatment of illness reveals
several problems. One is the delay between performance
of the behaviour and reinforcement. For prophylactic use,
the delay could be weeks or months. For therapeutic use,
acute conditions generally take days to noticeably im-
prove, even with modern drugs. Innumerable behavioural
responses would occur between use of a herbal product
and its reinforcement, from the prevention or reduction of
pain or discomfort, days or weeks later.
The occurrence of continuous reinforcement, which is
required for efficient acquisition of learned behaviour,
presents a problem with variability in the active ingre-
dients, and hence their effectiveness on the condition
being prevented or treated. In fact, variability in active
982 ANIMAL BEHAVIOUR, 70, 5
ingredients is one of the primary criticisms levelled
against medicinal herbs (Ernst 1998). Various medical
conditions would also be expected to respond differently
to the same medicinal herbs, lending additional inconsis-
tency to the reinforcement effects. Thus, the reinforce-
ment following the initial use of a plant part is likely to
be intermittent rather than continuous and not conducive
to learning. Finally, the necessity of multiple trials points
to a difficulty in the therapeutic use for acute illnesses.
Even if a herbal substance had reliable reinforcing proper-
ties, the infrequency of the illness would provide few op-
portunities for individuals to repeatedly experience the
reinforcing effects of a particular medicinal herb.
One possible model with regard to the operant learning
paradigm derives from research into mechanisms by
which animals with a deficiency in a nutrient acquire
a specific appetite for substances containing that nutrient.
A number of specific appetites are best explained on the
basis of postingestional reinforcing effects (Rogers &
Harper 1970; Zahorik 1977; Mori et al. 1991; Markison
et al. 1999). Representative of these studies is one on
rats made deficient in two essential amino acids (Markison
et al. 1999). Subjects showed no immediate tendency to
consume these amino acids in a liquid, but over a period
of 23 h they did gradually increase their intake of these
amino acids, presumably as a result of the postingestional
reinforcing effects. Reinforcement seemed to occur within
1–4 h and animals were reinforced continuously for
choosing the correct amino acid in multiple trials. Appli-
cation of this model, with a much shorter latency to alle-
viation of visceral discomfort than in the treatment of
illness and in the many trials with continuous reinforce-
ment, does not logically apply to the acquisition of herbal
medicine.
The possibility of a type of specialized learning in-
volving a biological predisposition should be mentioned.
The paradigm of conditioned food aversions, in which
a learned aversion occurs with only one experience with
an ingested toxin that causes gastrointestinal illness hours
later (Gustavson 1977), may suggest a sort of reverse para-
digm of one-trial learning with an ingested medicinal herb
that results in alleviation of illness or discomfort days later.
Aside from the convincing argument that learned food
aversions do not seem to greatly influence food choice
of animals in nature (Zahorik & Houpt 1981), herbal med-
icine differs in essential ways from conditioned food aver-
sions. Food aversions involve potential dietary items that
cause an onset of illness within a day or so. Medicinal
herbs are generally unpalatable and so they are not poten-
tial dietary prospects, and they may or may not lead to
gradual alleviation of illness over several days. At present,
no type of learning involving biological predispositions
appears to account for the origin of most types of herbal
medicine.
Natural Selection as a Basis for Herbal
Medicine
No learning paradigm seems to explain the use of herbal
medicines for the prevention or treatment of infections
and organ system dysfunction, but this use can be
explained by natural selection. The evolutionary explana-
tion suggests that the initial behavioural predisposition to
consume or use a herbal product was primarily an un-
learned response, selected because it enhanced survival
and/or reproductive success by reducing the likelihood or
severity of an illness or inflammatory process.
With regard to the prophylactic use of plant materials,
the evolutionary model implies that individuals that used
herbs that prevented illness when they were at risk were
less likely to experience loss of fitness than nonusers. For
example, wood rats that use larvacidal bay leaves in their
stickhouses (Hemmes et al. 2002) when they are at risk
should leave more healthy offspring than those not
engaging in the behaviour.
A useful model regarding the treatment of acute illness
concerns the one specific appetite for a micronutrient that
does not depend on postingestional learning. The specific
appetite for sodium is immediate in deficient animals,
requiring no experience with the altered state of deficien-
cy, and is considered innate (Nachman 1962; Handal
1965). Sodium deficiency is fairly common in nature
and sodium-deficient animals may be thought of as ill.
They will seek out and immediately ingest sodium-con-
taining substances with no experience with the deficiency.
The behaviour appears to be solely taste-guided, in that it
occurs even under experimental tests that eliminate the
influence of postingestive reinforcement (Markison et al.
1995). The sodium appetite model provides an analogue
for the seeking and consuming of medicinal herbs by indi-
viduals with an altered physiological state caused by dis-
ease or organ system dysfunction.
If an individual suffering from an inflammatory process
or a pathogen-induced illness seeks out and consumes
a plant part, the evolutionary explanation implies that the
herb has antimicrobial, anti-inflammatory, immunomod-
ulatory and/or analgesic effects, and that the individual
consuming such plants does not die or recovers more
quickly than do similarly sick individuals without this
predisposition. If this predisposition has a heritable basis,
then the selected behavioural predisposition should ex-
pand in the population.
Plant Physical or Chemosensory Markers
In addition to the requirement that putative medicinal
herbs actually be effective against various ailments and
enhance fitness, a requirement for the evolutionary
explanation for their use is that the herbal product has
a property that serves as a marker of some probability of
efficacy. Such a marker would be analogous to the salty
taste ‘marker’ of sodium-containing substances in the
sodium appetite model. In most instances of prophylactic
use of medicinal herbs, such markers would have to be
specific to the syndrome being prevented. Thus, a rough
surface may be a marker for leaves to be swallowed whole
(Huffman et al. 1996) and a type of volatile may be a marker
for plants used for nest fumigation (Hemmes et al.
2002).
 REVIEW 983

Many active ingredients of medicinal herbs are second- Table 3. Summary of medicinal and taste characteristics of 25 me-
ary plant compounds that serve as toxins for bacterial dicinal herbs
pathogens, parasites or insects that invade plants (Free- Characteristic Number of herbs (%)
land & Janzen 1974; Swain 1977; Deans & Ritchie 1987;
Knobloch et al. 1989; Langenheim 1994). The secondary
Medicinal effects
compounds, which are often toxic to mammalian species, Anti-inflammatory* 19 (76%)
generally have a repugnant, astringent or bitter taste, Antimicrobialy 14 (56%)
which may be considered a marker of toxicity (Garcia & Immunomodulatoryz 11 (44%)
Hankins 1975). Mammals in general dislike or reject bitter Analgesicx 7 (28%)
substances (Mattes 1985; Rouseff 1990; Glendinning Multiple medicinal effects
1994), and one would not expect plant parts to be food Two or more documented effects 24 (96%)
Anti-inflammatory and antimicrobial 10 (40%)
staples if they were strongly bitter or otherwise objection-
Anti-inflammatory and 6 (24%)
able-tasting, with high concentrations of secondary immunomodulatory
compounds. Anti-inflammatory and analgesic 7 (28%)
A bitter, astringent or repugnant taste, signalling un- Antimicrobial and immunomodulatory 7 (28%)
palatability of the plant part, is a logical marker of Taste characteristics**
medicinal efficacy. Of the 25 medicinal herbs featured in Bitter 10/22 (45%)
Table 2, I found information for 22 of them about the taste Astringent 6/22 (27%)
Bitter or astringent 16/22 (73%)
as they grow in nature, and a bitter, or astringent taste was Spicy or pungent 4/22 (18%)
mentioned for 16 of these (Table 3). Typical of what one Sweet 2/22 (9%)
might expect in traditional herbal medicine, interviews
with traditional healers in a Mixe Indian community re- *Anti-inflammatory effects through one or more recognized
vealed that bitter and astringent tastes were profiled in se- pathways.
lecting plants for medicinal purposes (Heinrich et al. yIncludes effects against pathogenic viruses, bacteria, fungi and pro-
1992). Bitter compounds have been used since antiquity tozoan parasites.
zBoost humoral or tissue immunity through one or more recognized
to treat illness, and even today are associated with medi- pathways.
cines and pharmaceutical efficacy (Brieskorn 1990). Even xEffects may stem from reduction of inflammation or recognized
the phrase, ‘a bitter pill’, conveys the notion that medi- pathway aside from inflammation. All analgesic herbs also have
cines are bitter. anti-inflammatory effects.
**Taste of plant part used as found in nature was available for only 22
Consistent with the evolutionary explanation, I suggest of the medicinal herbs.
that bitter or astringent plant parts were the focus of
natural selection for medicinal herbs used for therapeutic
purposes. The use of sweet tasting plant parts with
or pungent substances lies in the explanation of the
medicinal effects (e.g. licorice, ginseng) may have come
lowered threshold for rejection of bitter substances by
into use later as aspects of herbal medicine came under the
women in the first trimester of pregnancy. Profet (1988,
control of learning and social transmission.
1992) argued that this reduced acceptance of bitter foods
There are notable exceptions to the universal rejection
occurs in women when the developing fetus is most sus-
of bitter tasting substances. The sampling of bitter sub-
ceptible to the toxic ingredients of foods with bitter tastes,
stances with potentially medicinally active substances has
such as coffee and garlic. An illness-related, increased pref-
been noted repeatedly in mice (Glendinning 1994; Vitaz-
erence for bitter herbs, would be the reverse phenomenon.
kova et al. 2001) and has been reported in other mammals
Limited observations offer some evidence for increased
(Freeland & Janzen 1974). Modern humans are commonly
acceptability or attraction to bitter substances in sick
attracted to limited consumption of foods with a bitter
individuals. In her early studies, Goodall attempted to
taste, including quinine water, citrus fruits, coffee, tea
help sick chimpanzees at Gombe by offering bitter anti-
and cruciferous vegetables (Johns 1990; Rouseff 1990), al-
biotics in bananas; she reported that the sick chimpanzees
though the bitter ingredients are usually present in small
readily consumed the treated bitter-tasting bananas, but
amounts. These observations suggest that early humans
healthy chimps avoided them and waited for her to put
and animals in nature had a tendency to sample plant
out unadulterated bananas (J. Goodall, personal commu-
parts in their home range and knew what plant parts
nication, cited by Koshimizu et al. 1994). This anecdote is
were bitter, astringent or repugnant. While such behav-
consistent with the common belief that ill humans are tol-
iour is useful in avoiding toxic food staples, information
erant of bitter medicine, but this tolerance is not seen in
about bitter or astringent plant parts would also be avail-
healthy people or as sick individuals recover (Koshimizu
able to animals and people seeking bitter or astringent
et al. 1994).
plant parts when ill.
For an individual sick with an infectious disease, the
attraction to medicinal plant products brings up the Internal Stimuli Evoking Use of Medicinal
possibility that the illness itself results in a physiological Herbs by Sick Individuals
attenuation of the unpalatability of bitter-tasting plant
parts. That is, plant parts that would normally be rejected A requirement for the evolutionary explanation for
may be less unattractive to a sick individual. A precedent using herbal medicine to treat illnesses is the existence
for the physiological attenuation of a threshold for bitter of one or more physiological stimuli associated with the
984 ANIMAL BEHAVIOUR, 70, 5
illness that would provoke a predisposition to seek out
and consume an appropriate herbal product. For inflam-
matory conditions, the onset or exacerbation of pain or
discomfort would be the stimulus. For individuals with an
infectious bacterial or viral illness, the physiological
stimuli are likely to be not only pain or discomfort, but
also nausea, depression and fever, which are produced by
the endogenous pyrogens associated with the onset of
illness. The behavioural signs, currently referred to as
sickness behaviour, are relatively generic and occur re-
gardless of the type of illness (Hart 1988).
Social Transmission
Although I have argued that most uses of herbal
medicines, whether for treatment or prevention of illness,
reflected an evolved predisposition, social transmission of
the behaviour would increase the efficiency with which
appropriate plant parts would be located or used within
a group (Huffman & Hirata 2003, 2004). The best model
for such social transmission of herbal medicine is the facil-
itation of acceptance of novel food items by animals when
they observe another animal (the demonstrator) consum-
ing the food item (Galef 1996, 2002). An analogous exam-
ple would be social transmission of the consumption of
herbal products for prophylactic purposes, where the
products are consumed on a regular basis by healthy
animals.
The transmission of herbal medicine for therapeutic
purposes from individuals who have acquired the behav-
iour in the face of illness, to other group members who are
sick, would be expected in humans through verbal and
nonverbal communication. In light of recent research
revealing similar brain activation (anterior insula and
anterior cingular cortex) in people who observe others
suffering from painful stimuli (Carr et al. 2003; Singer
et al. 2004), one can envision a mentor ‘feeling another’s
pain’ and pointing to a bitter-tasting plant part that the
suffering individual should consume. For social transmis-
sion to occur involving the therapeutic use in animals,
imagine two animals that are sick at the same time, where
the demonstrator, who had already acquired the behav-
iour, consumes small amounts of otherwise unpalatable
plant parts in the presence of the similarly ill observer.
This type of social transmission, which requires that the
demonstrator be sick at the same time as the observer,
would be unlikely except in epidemic-like illnesses.
IMPLICATIONS OF THE EVOLUTIONARY
EXPLANATION
The hypothesis that herbal medicine arose through
natural selection raises a number of issues that merit
further discussion.
So Many Medicinal Herbs, So Many Maladies
A dilemma in understanding the origin of the thera-
peutic use of medicinal herbs is that it applies to the use of
herbal products for a wide array of diseases, each with
different etiologies, organ system involvements, time
courses and response characteristics. There are hundreds
of herbal products, and each may be recommended for
a list of maladies. Typical of such lists are the conditions
for which the medicinal herb barberry, Berberis vulgaris, is
indicated: amoebic dysentery, cholera, gastrointestinal in-
fections, hepatitis, eczema, psoriasis, limited bile flow and
gall bladder pains (Chevallier 2000, page 177). Based on
the specialized uses expected from modern synthetic
drugs, such a list evokes scepticism about efficacy of any
type. The medicinal herbs to which sick animals or hu-
mans may be attracted also undoubtedly differ in the de-
gree to which they contain a therapeutic agent
appropriate to the condition of the individual consuming
the herb.
Within the evolutionary perspective, then, what are
some responses to this dilemma? Of the sample of 25
medicinal herbs in Table 2, 76% (19) have anti-inflamma-
tory, 56% (14) antimicrobial, 44% (11) immunomodulatory
and 28% (7) analgesic effects, with 96% (24) having more
than one of these therapeutic effects (Table 3). Thus, as an
initial response to this dilemma, a medicinal herb is likely
to contain antimicrobial and/or immunomodulatory
agents as well as anti-inflammatory ones. The broad spec-
trum of medicinally effective agents in a single herb would
make the herb useful for a variety of maladies. Second, an-
imals and ancestral humans may have sought out more
than one bitter or astringent plant part. Third, although
a medicinal ingredient may not be an ideal match for a dis-
ease syndrome, one or more constituents might provide
some benefit. For example, one would not ordinarily treat
an upper respiratory viral disease with an antibiotic, but in
nature a herbal antibiotic could reduce secondary bacterial
complications. An anti-inflammatory agent may allow an
individual with a painful condition brought about by
a bacterial pathogen to go about providing care of depen-
dent offspring sooner, even though an antimicrobial
agent is medically indicated.
Ineffective and Toxic Medicinal Herbs
The evolutionary explanation must account for not
only the vast array of herbal plants and variety of illnesses,
but also the use of plants that share the chemosensory
markers of medicinally effective herbs but that either have
no medicinal effects or may be toxic. These criticisms are
commonly levelled against putative medicinal herbs
(Ernst 1998). At the outset, one would expect the practice
of herbal medicine to include plants having a level of effi-
cacy distributed along a continuum ranging from effective
to ineffective. Medicinal herbs, even with a fraction of the
potency or safety of modern drugs, could be the basis for
the selection of their use as long as some benefit in fitness
accrued to the user.
There are two aspects to the consideration of ineffective
medicinal herbs. One involves plants that are not effective
for some disease syndromes but are effective for other
syndromes. Thus, a herb may appear to be worthless if
tested for antibacterial effects but may alleviate a painful

flare-up in an arthritic knee joint. Unless the herb is tested
for several types of effects, its value may remain unknown.
The other aspect involves plants that have no medicinal
value whatsoever. Both aspects are addressed in the same
way. As with other behavioural traits related to fitness,
a behaviour need only have an occasional effect on fitness
to be maintained by natural selection. This prediction
should be especially true of behaviours that influence
survival of the individual in question or welfare of
offspring. The continued use of some ineffective herbs
that are indistinguishable from effective herbs would be
predicted by the evolutionary explanation.
Another prediction from the evolutionary explanation
is that herbs that are medicinally effective, but toxic to
one or more organ systems, may be retained in the herbal
medicine repertoire. Although the intention is to avoid
such side effects with modern medicines, the benefits
could outweigh the costs of some loss of organ system
tissue for animals and humans living in nature, and at
a life-or-death juncture, or if the care of dependent young
is threatened. This is the commonly accepted paradigm
for the evolutionary persistence of the adaptive value of
the microbe-suppressing fever response of sick animals
when febrile temperatures reach tissue-damaging levels
(Ewald 1980; Hart 1988).
Advent of the Placebo Effect
Human religious practices that use putative medicinal
herbs are rife with verbally communicated expectations of
recovery or improvement. This expectation, and the sub-
sequent influence on recovery or improvement, is known
as the placebo effect (Shapiro & Shapiro 1997). The place-
bo effect would, of course, apply to herbal products that
are both medicinally effective and ineffective. Clearly
the placebo effect is enhanced when a patient is told by
a medical authority to expect to improve, regardless of
whether the authority is a modern physician or a tribal
healer. The fact that a proportion of patients improve by
virtue of the placebo effect would help to maintain the
use of an ineffective herbal product. The placebo effect ap-
pears to be a function of human cultural and linguistic
properties and not generally available to nonverbal
animals.
CONCLUSIONS: ANIMAL ANALOGUES
OF MEDICINE
I have argued that the origin of herbal medicine in
animals and ancestral humans was largely a reflection of
natural selection stemming from fitness benefits to indi-
viduals that engaged in this behaviour. The cornerstone of
this explanation, as applied to therapeutic use, is that
individuals suffering from acute illness or inflammatory
conditions are drawn to bitter or astringent plant parts
that are likely to have anti-inflammatory, antimicrobial,
immunomodulatory and/or analgesic effects that may be
effective in a nonspecific manner. This perspective pro-
vides an explanation of the wide array of putative
medicinal herbs that are suggested for numerous and
REVIEW 985
varying maladies. The evolutionary perspective explains
why ineffective or toxic plant parts may be maintained in
the repertoire of putative medicinal herbs. The evolution-
ary perspective also provides an explanation for fitness
benefits that may be very delayed and indefinite, a per-
spective relevant to both the prophylactic and therapeutic
types of herbal medicine.
Although the analogues of human herbal medicine (and
subsequently pharmaceuticals) arguably occur in non-
human animals, the field of human medicine encom-
passes more than just using medicinal substances to
prevent or facilitate recovery from disease. We recognize
as essential aspects of human medicine the value of
nursing and caring for sick group members, immunizing
young and vulnerable group members against agents of
infectious disease, and the removal or isolation (quaran-
tine) of sick individuals that place other group members at
risk. In a previous review, I identified several behavioural
strategies that are analogues of these essential aspects of
human medicine that nonhuman animals use to deal with
pathogens and parasites (Hart 1990). As with my argu-
ment that learning is an unlikely explanation for most as-
pects of the acquisition of the practice of herbal medicine,
learning is also an unlikely explanation of these other
strategies of disease control, which may be viewed as pil-
lars of modern medicine with analogues in animal behav-
iour. Correspondingly, it is unlikely that ancient humans
learned about aspects of medicine from nonhuman ani-
mals, but rather that the same natural selective forces in-
volved in combating pathogens and parasites led to
similar behaviour defence strategies in both nonhuman
animals and ancient humans.
Acknowledgments
Research on wood rats, which provoked my thinking
about some of the concepts in this review, was supported
by grant IBN-96-17407 from the National Science Foun-
dation. The author thanks colleagues Lynette Hart, Dick
Coss, Dorothy Geitzen, Mike Mooring and the late Dale
Lott for valuable input and the constructive criticisms of
four anonymous referees.
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