FORUM
Committed Phloem Ingestion of Aphids (Homoptera: Aphididae):
Difficulties in Its Application to Host-Plant Resistance Studies
MELAKU GIRMA,l GERALD E. WILDE,l JOHN C. REESE,l
AND ELAINE A. BACKUS2
Environ.Entomol. 23(5): 1061-1065 (1994)
ABSTRACT The application of the concept of committed phloem ingestion (CPI) of
aphids was studied for different aphid species feeding on resistant and susceptible plants.
Greenbugs, Schizaphis graminum (Rondani) (biotype E), were monitored electronically
on wheat, Triticum aestivum L., and Sorghum bicolor (L.) Moench, cultivars, and Russian
wheat aphids, Diuraphis noxia Mordvilko, were monitored on wheat cultivars. The dura-
tion of phloem ingestion (PI) events of these aphids was not randomly distributed and
"'53% of them were <60 min in duration. The minimum PI duration to be considered as
committed feeding (threshold of CPI) was determined for each aphid-host combination.
Thresholds varied depending on the aphid species and host plants, indicating the difficulty
ofassigning a single threshold value that could be useful as a measure of host acceptability.
Also, if a given threshold duration is chosen as a criterion for having achieved CPI (e.g., 15
min), the proportion of PI events that continues for a long time period varies from host to
host. Therefore, we recommend that use of this parameter be discontinued, and that
measurements ofthe time from the beginning of the experiment to PI and the time from the
initiation of a probe to PI be substituted for time to first CPI.
KEY WORDS aphids, probing behavior, electronic monitor
COMMITTEDPHLOEMINGESTION(CPI) has been
a widely used variable in the quantification of
feeding behavior of aphids. It was first intro-
duced by Montllor et al. (1983). They chose 15
min of PI as a threshold for an aphid commit-
ment to long-term feeding because "an aphid
which ingested from the phloem continuously
for at least 15 min tended to remain feeding in
phloem for up to several hours." Subsequently,
this threshold has been used to estimate other
feeding parameters such as time to the first CPI
(Montllor et al. 1983, Ryan et al. 1987) and po-
tential phloem ingestion index (PPII) (Montllor
et al. 1983, Girma et al. 1992) as measures of host
acceptability. Unfortunately, publications have
not always made it clear whether CPI was mea-
sured from the beginning of the experiment or
the beginning of the probe, also known as pene-
tration. Time to the first CPI refers to the time
taken for an insect to reach a phloem sieve ele-
ment and ingest its sap for at least 15 min. Time
to the first x-wave, or phloem contact, measures
only the relative difficulty in penetrating plant
tissue en route to the phloem. However, time to
the first CPI includes both plant morphological
and physiological factors that affect aphids be-
1 DepartmentofEntomology, KansasStateUniversity,Man-
hattan,KS66506-4004.
2 Department of Entomology, University of Missouri-
Columbia,Columbia,MO65211.
fore they reach the phloem as well as sap quality
while ingesting from the phloem.
The CPI parameter has been used widely with
greenbugs, Schizaphis graminum (Rondani),
feeding on sorghum, Sorghum bicolor (L.)
Moench (MontHor 1985, MontHor et aI. 1983); on
wheat, Triticum aestivum L. (Niassy et al. 1987,
Ryan et al. 1987); on barley, Hordeum vulgare L.
(Peters et al. 1988); on corn, Zea mays L. (Mc-
Cauley et al. 1990) and on oats, Avena sativa L.
(MontlIor & Gildow 1986). It also has been used
in feeding behavior studies of other aphids, in-
cluding the hop aphid, Phorodon humuli
(Schrank) (Dorschner & Baird 1989); Russian
wheat aphid, Diuraphis noxia Mordvilko (Girma
et al. 1992); and bird cherry-oat aphid, Rhopal-
osiphum padi (L.) (Montllor & Gildow 1986).
Although CPI has been used by many research-
ers, its 15-min threshold value has not been
quantified using experimental data. MontUor
(1991) and Backus (1994) have suggested several
interpretational problems with the concept.
However, until our work, no one has attempted
to test CPI quantitatively. Dorschner & Baird
(1989) assigned a posteriori a 60-min CPI thresh-
old to the ingestion of the hop aphid. Studies by
Girma et al. (1992) and Montllor (1991) showed'
that the threshold for CPI can vary among aphid
species and even among individuals within spe-
cies as they feed on different hosts. The actual
threshold value researchers use to determine
0046-225X19411061-1065$02.00/0 © 1994 Entomological SocietyofAmerica
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