AQUATIC MICROBIAL ECOLOGY
Aquat Microb Ecol
Vol. 44: 85–96, 2006 Published August 16
INTRODUCTION
Studies in which phagotrophy is addressed are still
seriously hampered by tedious and often time-consum-
ing methods (Riemann et al. 1995). Over the last
decades, many methodologies have been developed to
detect phagotrophy in heterotrophic and mixotrophic
phytoplankton species. Visually detected and manu-
ally counted tracer-based techniques are the most
common approaches for estimating phagotrophy in
algae (Havskum & Riemann 1996, Granéli & Carlsson
1998, Legrand et al. 1998, 2001, Pålsson 2001). Surro-
gate prey, including dye and starch particles, fluores-
cent microspheres (Smalley et al. 1999), fluorescently
labeled bacteria and algae have been used to investi-
gate feeding behaviors and to measure ingestion rates
of planktonic protists (Li et al. 1996).
Although the use of fluorescently labeled prey has
resulted in an increased understanding of phagotro-
phy, advances in this field have been slow (Granéli &
Carlsson 1998). Usually a large number of cells must
be counted manually with the help of a microscope in
order to achieve statistically significant ingestion rates,
a monotonous and lengthy procedure. In addition, epi-
fluorescence microscopy has a low resolution with
regard to detection of ingested fluorescent prey, espe-
© Inter-Research 2006 · www.int-res.com *Email: wanderson.carvalho@hik.se
Acidotropic probes and flow cytometry: a powerful
combination for detecting phagotrophy in
mixotrophic and heterotrophic protists
Wanderson F. Carvalho*, Edna Granéli
Marine Science Department, University of Kalmar, 391 82 Kalmar, Sweden
ABSTRACT: Studies with phagotrophic organisms are hampered by a series of methodological con-
straints. To overcome problems related to the detection and enumeration of mixotrophic and hetero-
trophic cells containing food vacuoles, we combined flow cytometry and an acidotropic blue probe as
an alternative method. Flow cytometry allows the analysis of thousands of cells per minute with high
sensitivity to the autofluorescence of different groups of cells and to probe fluorescence. The method
was first tested in a grazing experiment where the heterotrophic dinoflagellate Oxyrrhis marina fed
on Rhodomonas salina. The maximum ingestion rate of O. marina was 1.7 prey ind.
–1
h
–1
, and the fre-
quency of cells with R. salina in the food vacuoles increased from 0 to 2.4 ± 0.5 × 10
3
cells ml
–1
within
6 h. The blue probe stained 100% of O. marina cells that had R. salina in the food vacuoles. The
acidotropic blue probe was also effective in staining food vacuoles in the mixotrophic dinoflagellate
Dinophysis norvegica. We observed that 75% of the D. norvegica population in the aphotic zone pos-
sessed food vacuoles. Overall, in cells without food vacuoles, blue fluorescence was as low as in cells
that were kept probe free. Blue fluorescence in O. marina cells with food vacuoles was 6-fold higher
than in those without food vacuoles (20 ± 4 and 3 ± 0 relative blue fluorescence cell
–1
, respectively),
while in D. norvegica cells were 4.5-fold brighter than the ones without food vacuoles (291 ± 155 and
64 ± 23 relative blue fluorescence cell
–1
, respectively). The use of acidotropic probes can prevent fix-
ation artifacts such as regurgitation of food vacuoles and changes in the cellular characteristics. The
combination of flow cytometry and an acidotropic probe proved to be an efficient tool in detecting
phagotrophy in mixotrophic and heterotrophic marine phytoplankton species.
KEY WORDS: Mixotrophy · Phagotrophy · Acidotropic probes · Flow cytometry
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