The ability to respond to chemical stimuli is among
the key sensory functions found in almost all living organ-
isms. Green flagellate algae including Chlamydomonas
that are model objects in many fields of biological
research use chemical stimuli to adapt to their environ-
ment by specific chemoinduced behavior. Although the
first observations of such responses in Chlamydomonas
date back to the 19th century [1], our knowledge of them
is still at the level of phenomenology. On the contrary,
significant progress has been achieved in investigation of
chemoregulation of behavior in other cells. Best of all has
been characterized chemotaxis in motile cells in humans
and animals [2, 3], as well as chemotaxis in some
prokaryotes (mainly enterobacteria [4, 5]). Also, much is
known about chemotaxis in the lower eukaryotes: the
myxamoeba Dictyostelium [6, 7] and the ciliate
Paramecium [8, 9].
With the exception of Paramecium, success in inves-
tigation of chemoregulation of motility in these particular
objects was due to their being amenable for molecular
genetic analyses. Recently completed sequencing of the
whole nuclear genome of Chlamydomonas reinhardtii [10]
created a possibility for rapid progress in research into
chemoreception in this microorganism as well. This opti-
mism is supported with the remarkable progress in inves-
tigation of other physiological functions of this alga
achieved since it became the object for establishment of
nucleotide sequence databases. In particular, this facilitat-
ed the identification of rhodopsin receptors for phototaxis
and the photophobic response in C. reinhardtii [11, 12].
The nuclear genome of C. reinhardtii contains 100 to
110 million base pairs combined into 17 linkage groups
(chromosomes). Its sequencing has been undertaken by
the US Department of Energy Joint Genome Institute.
By the time of this review, initial assembly of the infor-
mation obtained by sequencing overlapping short frag-
ments of the genome has been completed. The results of
this work, as well as their preliminary analysis, can be
accessed at the JGI web page: http://genome.jgi-
psf.org/chlre2/chlre2.home. html.
One of the reasons for Paramecium being a popular
experimental organism is that it is suitable for electro-
physiological measurements. It has been found that
mechanisms for chemical signaling in Paramecium
involve bioelectrical processes in the cell membrane, and
detailed studies on these processes have been carried out
[9]. Although Chlamydomonas is too small for reliable
microelectrode recording widely used in Paramecium,
methods for extracellular measurement of asymmetrical-
ly localized transmembrane currents in this and related
algae have been developed [13-15]. It has been shown that
photoexcitation of rhodopsins leads to generation of such
currents, which play a key role in photoregulation of
motility in these microorganisms [16]. The possibility of
photoelectric recording prompted investigation of pho-
toreception and photomovement in Chlamydomonas [17].
Biochemistry (Moscow), Vol. 70, No. 7, 2005, pp. 717-725. Translated from Biokhimiya, Vol. 70, No. 7, 2005, pp. 869-879.
Original Russian Text Copyright © 2005 by Govorunova, Sineshchekov.
REVIEW
0006-2979/05/7007-0717 ©2005 Pleiades Publishing, Inc.
* To whom correspondence should be addressed.
Chemotaxis in the Green Flagellate Alga Chlamydomonas
E. G. Govorunova* and O. A. Sineshchekov
Faculty of Biology, Lomonosov Moscow State University, 119992 Moscow, Russia;
fax: (7-095) 939-4309; E-mail: egovoru@yahoo.com
Received July 14, 2004
Revision received August 12, 2004
Abstract—Behavior of the green flagellate alga Chlamydomonas changes in response to a number of chemical stimuli. Specific
sensitivity of the cells to different substances might appear only at certain stages of the life cycle. The heterogamous species
C. allensworthii demonstrates chemotaxis of male gametes towards pheromones excreted by female gametes. In C. reinhardtii
chemotaxis towards tryptone occurs only in gametes, whereas chemotaxis towards ammonium, on the contrary, only in veg-
etative cells. Chemotaxis to different chemical stimuli might involve different mechanisms of reception and signal transduc-
tion, elucidation of which has only recently begun. Indirect evidences show that the cells likely respond to tryptone with
changes in the membrane electrical conductance. The recently completed project of sequencing the whole nuclear genome
of C. reinhardtii provides the basis for future identification of molecular elements of the chemosensory cascade in this alga.
Key words: chemoreception, signal transduction, photoreceptor current, ammonium, tryptone, gametogenesis