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Heliozoa PDF
Heliozoa PDF
Introduction
Heliozoa (Gk helios sun, zoon animal) are unicellular involved in many physiological functions, including
phagotrophic animalcules with radiating axopods like the recognition of environmental stimuli, cell contraction,
rays of the sun. These eukaryotes belong to the kingdom locomotion and food capture. Axopods are stiened by a
Protozoa and the supertaxon Actinopoda. They live at rigid, though labile, axial rod which consists of a complex
shallow depths in freshwater, brackish or marine environ- assembly of parallel microtubules and crosslinking
ments. Most species are free-living, oating just above bridges. The microtubule-based cytoskeleton is seen to be
solid substrates or rolling among aquatic plants. They are organized in specic and complex patterns when viewed in
never found in turbid water, which produces retraction of cross-section; for instance, including repeating motifs of
cytoplasmic extensions used for feeding. Reproduction is double interlocking coils, elongated hexagons, regular
generally asexual but sexual processes with production of hexagons and equilateral triangles, as well as other more
complicated gures.
amoeboid or agellate gametes have been described.
One of the distinctive features of the axopods is their
sensitivity to a variety of physical stimuli, such as
mechanical or electrical shocks, ultrasound, cold, high
Description and Characteristics pressure and chemical agents. In some species, mechanical
shocks induce slow membrane depolarization. In other
Most species of Heliozoa are spherical, measuring 0.05 species mechanical or electrical stimulation elicits a
0.5 mm in diameter, although some are more complex, rapid, high-amplitude action potential; while in a few
comprising a cytoplasmic base bearing a contractile stalk cases, the cell membrane is mechanically insensitive,
and an oval head. The cell membrane is covered with a stimulation up to 300 shakes per minute for 5 minutes
mucous coat that can include dispersed mineral or organic producing no obvious eect. Chemotactic behaviour
scales and spicules. The cytoplasm of heliozoans is never has been demonstrated using prey organisms or
enclosed in a central capsule as it is in radiolarians. Long, particles coated with specic proteins, suggesting chemo-
granule-studded processes known as axopods radiate from sensitivity. Another distinctive feature of heliozoan
the cell body. Axopods (and the stalk when present) are axopods is their remarkable contractility. The highly
strengthened by highly ordered bundles of unstable ordered microtubular arrays responsible for main-
microtubules that radiate from one central mass of taining the radial shape can dissociate almost completely
brillar-granular material in the cell body, or from within a few milliseconds in response to external
multiple sites associated with the nuclear envelope. The stimuli. During rapid contraction, the microtubular rods
cytoplasm includes microtubule organizing centres rst fragment, perhaps by the action of microtubule-
(MTOCs), Golgi elements and endoplasmic reticulum, associated severing proteins, then disassemble from the
mitochondria with tubular cristae, one or several nuclei ends of fragments. Axopodial arrays then regrow from
and, in some species, endosymbiotic algae. Prey organisms, the MTOC. Certain chemicals, including microtubule
extrusive organelles involved in prey capture, food inhibitors, urea and heavy and light metal ions, as
vacuoles, lysosomes and lipid droplets are present in the well as mechanical and electrical stimuli can induce
supercial cytoplasm (Figure 1). microtubule destabilization and axopodial retraction.
Only agents that stabilize microtubules, such as heavy
Axopods water, taxol (a drug isolated from the yew tree Taxus
brevifolia) or calcium-free media are able to stabilize this
Axopods are slender extensions that project 0.040.8 mm highly unstable microtubule-based array under experi-
from the cell body into the surrounding water. They are mental stimulation.
ENCYCLOPEDIA OF LIFE SCIENCES 2002, John Wiley & Sons, Ltd. www.els.net 1
Heliozoa
Figure 1 Structure of two heliozoa: a centrohelid heliozoon (left) and the actinophryid species Actinophrys sol (right).
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This composite layer is periodically cast o and replaced by ence. In the stalked species, A. contractilis, the life cycle
a new one. comprises two phases, asexual multiplication through
binary or multiple ssion and sexual reproduction, the
whole heliozoan giving rise to thousands of biagellated
gametes that are shed synchronously. Gametes swim
Behaviour actively, then are likely to fuse.
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Heliozoa