Attitudes towards the risk of damage by biological
control agents to plants other than the target weed, and the use of
host-specificity testing to minimise that risk, have changed through the
history of biological control of weeds.
In the early programs by Hawaii, no host-testing was
undertaken (Waterhouse and Norris
1987). Observations were made in the field in Mexico, and if the
insects were seen feeding only on lantana, it was assumed that they were
sufficiently host-restricted for safe introduction. Hawaii continued to
rely on field observations in the country of origin to determine
host-specificity, until at least the 1950s. The unique and restricted
flora of Hawaii, with few relationships to plants in other continents,
probably reduced any problem of attack on plants other than the target
weed. However, problems did arise when insects were introduced into other
countries on the basis of their use in Hawaii and without further tests.
For example, the lantana tingid Teleonemia scrupulosa
was introduced into East Africa in the 1960s without further testing,
where it caused problems by moving onto the crop sesame when the lantana
was defoliated. There was also attack on teak trees. Economic damage was
not great, but the attack caused problems and it is likely that the insect
would not have been introduced if host-testing had been carried out first.
Host-specificity testing was first used in Australia in
the 1920s, in the major campaign against the prickly pears (Dodd
1940). Field observations, where insects were seen to be feeding only
on Cactaceae in the wild, together with the known association of the
insect type with Cactaceae, were still seen as the chief proof of host
specificity. Host-testing was undertaken mainly to satisfy the general
public that crop plants would not be damaged. Initially, both choice tests
and no-choice tests were used, but the choice tests were quickly abandoned
in favour of no-choice tests. In choice tests, candidate agents are
confined for varying periods of time with the test plant species together
with the target weed. In no-choice tests, candidate agents are confined
usually with only one test plant species and without the target weed. The
tests used were starvation tests, that is, the insects were confined on
the test plant until they died or developed through to the next stage. As
the purpose of the tests was to prove that crop plants were safe, only
plants of economic importance were tested; there was no concern for native
Australian plants other than their economic value.
The conservative principle was adopted from the start (Dodd
1940): that is, an insect was rejected if it could complete
development on a test plant, even if oviposition on the plant would not
normally occur, and even where the insect was known not to attack the test
plant in the field in its native range. Similarly, if one species in a
genus was considered unsafe, the whole genus was rejected. Several stages
were tested; always newly-hatched larvae or nymphs, and usually half-grown
larvae and adult oviposition as well.
By the 1950s, the next major period of activity,
host-testing was seen as an essential part of a weed biological control
program. No-choice or sequential host-specificity tests of varying
duration became the accepted methodology for the determination of host
specificity until the late 1960s (Harris and Zwolfer
1968). Sequential tests involve the sequential presentation of a
series of test plants in a no-choice situation, with each plant species
usually exposed to the candidate agent for a relatively short time.
In the late 1960s Harley (1969)
advocated choice as opposed to no-choice tests. It was argued that choice
tests are a more 'natural test' of host range, as the target weed will be
usually be present in mixed stands with other plants in the field
situation (Cullen
1990). Choice tests may lead to fewer incidences of feeding on test
plant species and less rejection of 'safe' insects. Discussions on the
'best' methods for host-specificity testing continue, and a mixture of
choice and no-choice tests is usually used.
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