AMER. ZOOL., 35:174-185 (1995)
Alvinellids and Sulfides at Hydrothermal Vents of the
Eastern Pacific: A Review1
S. K I M JUNIPER AND PASCALE MARTINEU
Centre de Recherche en Giochimie Isotopique et Gtochronologie (GEOTOP), University du Quebec a Montreal,
P.O. Box 8888, Station A, Montreal (Quebec) H3C 3P8 Canada
SYNOPSIS. At Eastern Pacific hydrothermal vents, alvinellid polychaetes
are among the first metazoans to colonize newly formed surfaces of sulfide
chimneys. In this environment of rapid mineral precipitation, alvinellids
are confronted by steep physico-chemical gradients and high temporal
variability. This paper examines the interaction of alvinellids with chimney mineralization processes and then reviews what is known of mechanisms that could enable these worms to deal with potentially toxic levels
of sulfide in their environment. Studies of sulfide chimneys consistently
show mineralogy to be locally modified around alvinellid tubes. This may
be linked to sulfide oxidation products that accumulate in tube material
or to the circulation of seawater through the tube. At high worm densities,
these local effects may have a significant influence on larger scale sulfide
accretion processes that determine chimney morphology. Alvinellid polychaetes may have several lines of defense against sulfide. Tubes and
mucous layers could act as passive barriers to reduce inward diffusion of
sulfide across posterior surfaces. Colonization of epidermal and tube surfaces by bacteria that might be sulfide oxidizing, could create an active
external barrier in some species. Sulfide oxidation by tissue homogenates
has been demonstrated in two Paralvinella species, where it may serve to
protect oxidative respiration from sulfide entering worm tissues. Sulfide
binding in blood has not been studied in any of the alvinellids.
INTRODUCTION
the East Pacific Rise (EPR). The northeast
Since the discovery of black smoker Pacific palm worm Paralvinella palmiforchimneys in 1979 (Spiess et al, 1980), bio- mis and the EPR Paralvinella grasslei are
chemical and physiological studies of chim- found on cooler portions of chimneys and
ney organisms have primarily focused on are not restricted to the chimney habitat
prokaryotes. Specialized metazoan com- (Desbruyeres et al, 1985; Desbruyeres and
munities also exploit this extreme and rap- Laubier, 1986; Tunnicliffe et al, 1993).
idly changing environment (Desbruyeres et
Alvinellidae is a terrebellomorph polyal, 1985; Tunnicliffe and Juniper, 1990). chaete family that is endemic to hydrotherAmong the first animals to colonize newly mal vent habitats (Desbruyeres and Lauformed chimney surfaces are the alvinellid bier, 1986, 1991). According to Tunnicliffe
worms (Juniper et al, 1992). The northeast et al. (1993), the eastern Pacific alvinellid
Pacific vents are the habitat of the "sulfide worms, Alvinella spp. and Paralvinella spp.,
worm," Paralvinella sulfincola Desbruyeres likely evolved from a common ancestor that
and Laubier (Tunnicliffe et al, 1993), while colonized the East Pacific Rise and the Juan
the "Pompeii" worms Alvinella pompejana de Fuca Ridge before separation of the two
and A. caudata occupy a similar niche on ridge systems some 25 million years ago.
Alvinella spp. is the more derived form and
Desbruyeres and Laubier (1991) propose an
early separation of the genus Alvinella from
I
~ „
.
. ,„,._,
1
From The Symposium Life with Sulfide presented e v o lutionarv lines taken hv the Pnrnlvirtplln
atthe Annual Meeting of the American Society of Zool- ev <"Uiionary lines taken oy me Paralvinella
ogists, 27-30 December 1993 at Los Angeles, Califor- species. This review will be limited to two
nia.
species of Alvinella {A. pompejana and A.
174
175
ALVINELLIDS AND SULFIDES
caudata) from the EPR and two paralvinellids from the Juan de Fuca Ridge (P. sulfincola and P. palmiformis) (Fig. 1). We
examine the influence of alvinellid polychaetes on sulfide mineralization processes
and consider adaptive mechanisms that may
enable them to tolerate the high sulfide levels that are characteristic of the chimney
environment.
The sulfide chimney environment
The smoking chimneys and spires that
develop around high temperature vents are
the focal points of sulfide mineral precipitation on unsedimented spreading ridges
(Hekinian et al, 1983; Hekinian and Fouquet, 1985; Tivey and Delaney, 1986; Hannington and Scott, 1988). Chimney growth
begins when sea water, entrained into a jet
of high temperature (35O°C) fluid, is heated
to supersaturation with respect to CaSCy
This causes the precipitation of anhydrite
and the upward growth of primitive chimney walls in the form of a porous anhydrite
shell (Haymon and Kastner, 1981; Haymon, 1983). Induration of this shell by further precipitation of anhydrite and other
minerals eventually begins to isolate hydrothermal fluid within the structure from surrounding seawater to the point where significant physico-chemical gradients develop
across the chimney walls. This initiates the
second phase of chimney development.
While anhydrite continues to accrete on the
outer walls and on the distal portion of the
growing chimney, hydrothermal fluids
migrate outward through the walls, dissolving and replacing anhydrite with sulfide
minerals. At the same time, high-temperature conditions within the chimney conduit favor the inward growth of Cu-Fe rich
sulfides. Chimneys can grow in this manner
up to several meters in height, accreting new
anhydrite material around the vent orifice,
and later infilling and replacing it with sulfides. Instead of anhydrite, direct precipitation of barite or sphalerite can form the
early shell (Paradis et al, 1988). The subsequent processes are essentially the same
in all cases. Early walls are replaced and
sealed by outflowing hydrothermal fluid that
is precipitating sulfide.
150"
130"
110"
90" W
60'N
C-I15-S
FIG. 1. Map of Eastern Pacific spreading centers
including the East Pacific Rise (EPR) and the Juan de
Fuca Ridge. Alvinellid species discussed in this paper
are situated in relation to their respective seafloor
spreading centers.
Colonization of new chimney surfaces by
alvinellid polychaetes begins during the early
stages of secondary mineralization. It is difficult to define precisely the range of conditions that constitute the worms' habitat.
All physico-chemical habitat information
must be collected remotely from submersibles, and in this environment of sharp
physico-chemical gradients, reliable recording of conditions within the worm's immediate microenvironment is difficult. Temperatures of 20-80°C have been measured
on surfaces colonized by Paralvinella sulfincola and P. palmiformis (Juniper et al,
1992), and it is most likely that the worms
regularly experience temperatures within the
lower part of this range. Alvinella spp. is
frequently found at temperatures near 40°C
(Fustec et al, 1987) and has recently been
reported moving over a 105°C surface
(Chevaldonne et al, 1992). However, physiological and biochemical studies (reviewed
in Childress and Fisher, 1992) indicate that
Alvinella spp. are unlikely to be exposed to
temperatures above 40°C for extended periods. Tunnicliffe et al. (1993) emphasize the
importance of turbulence and likelihood of
176
S. KIM JUNIPER AND PASCALE MARTINEU
sw
FIG. 2. Spatio-temporal variability in the distribution
of Alvinella spp. on chimneys at 13°N EPR. Inactive
(mineralized) portions of chimneys have no Alvinella
spp. colonies. A) Distribution of Alvinella spp. colonies
on different faces of Parigo chimney, adapted from
Chevaldonne and Jollivet (1993). B) Temporal evolution of Alvinella spp. colonization on Actinoir chimney, in relation to chimney growth and mineralization.
Adapted from Fustec et al. (1987).
occasional blasts of hot water in the upper
part of the 20-80°C range.
Sulfide data for this microhabitat are rare.
Measurements with a flow injection scanner
indicate that northeast Pacific paralvinellids
commonly experience 200-300 MM sulfide
concentrations (Sarrazin et al, unpublished
data). Turbulent mixing and resulting temporal fluctuation of environmental conditions (Johnson et al, 1988a; Chevaldonne
et al., 1991; Childress and Fisher, 1992) are
probably essential to survival of these ani-
mals: even during periods of high sulfide
concentration, small amounts of dissolved
oxygen would still be periodically available
to the worms.
Colonization of chimneys by
alvinellid polychaetes
Alvinellids presumably inhabit the inhospitable sulfide chimney environment to have
access to abundant food. All four worms
discussed here ingest particles and are presently considered to rely on deposit feeding
for growth and energy metabolism (reviewed
by Juniper, 1994). No internal symbionts
have been found in any of these species.
Bacteria that grow on sulfide and sulfate
particles are the most likely food source,
although bacterial abundance and productivity on chimney surfaces have never been
systematically quantified (Juniper, 1994).
Alvinella pompejana and A. caudata colonize two types of high temperature edifices: black smoker chimneys where undiluted fluids reach temperatures near 350cC
and "snow ball" diffusers where 200-300°C
fluids flow through 1-2 m high mound-like
structures (Desbruyeres et al, 1983, 1985;
Desbruyeres and Laubier, 1991). There
appears to be considerable spatial and temporal variability in the colonization of black
smoker chimneys by Alvinella spp. Chevaldonne and Jollivet (1993) used videoscopic methods to illustrate the patchy distribution of Alvinella spp. habitat on a
chimney at 13°N EPR (Fig. 2a). An example
of temporal changes in the distribution of
Alvinella spp. habitat on a growing black
smoker chimney is presented in Figure 2b.
Although migration of adult Alvinella spp.
on individual chimneys has not yet been
documented, adults have been suggested to
leave their tubes, and colonize newly formed
chimney surfaces (Fustec et al, 1987). Alvinella spp. were first described to construct
U-shaped tubes whose open ended design
tube allowed convective cooling of the tube
interior by indrawn seawater (Desbruyeres
et al, 1985). While frequently discussed in
subsequent publications, the U-shaped
morphology has never been systematically
documented. Chevaldonne et al. (1991)
recently proposed that tube irrigation and
ALVINELLIDS AND SULFIDES
thermoregulation in Alvinella spp. are
accomplished behaviorally by movement in
and out of the tube. As well, Chevaldonne
and Jollivet (1993) state that Alvinella spp.
do not build U-shaped tubes.
The zone of early sulfide mineralization
on growing chimneys is the preferred habitat of the northeast Pacific "sulfide worm",
Paralvinella sulfincola (Juniper etai, 1988;
Tunnicliffe and Juniper, 1990) (Fig. 3). Prior
to worm colonization, a thickening of the
anhydrite walls of the chimney with concomitant decrease in temperature and sulfide levels appears be necessary (Juniper et
ai, 1992). The process of anhydrite shell
formation, secondary mineralization and
colonization by sulfide worms can apparently occur in a matter of days. A time lapse
camera study of the growth of small spires
on a sulfide mound at Axial Seamount (Fig.
4a) documented the colonization of newly
mineralized surfaces by adult P. sulfincola
over a 33 day period (Fig. 4b; Juniper et ah,
1992). The tube of P. sulfincola is a multilayered structure that is less organized at the
ultrastructural level than the tubes of A.
pompejana (Juniper, 1994). The outer surface of the tube remains soft and tends to
trap mineral particles (Juniper et ai, 1986).
The northeast Pacific "palm worm," Paralvinella palmiformis, is usually found in
association with vestimentifera, on cooler
portions of chimneys and at diffuse flow
vents on basalt surfaces (Tunnicliffe and
Juniper, 1990). Mucus secreted by the palm
worm, rather than being used in tube construction, is continuously sloughed off the
epidermal surface. This may protect the
worm from precipitating mineral particles,
and possibly serve as a barrier to chemical
diffusion (see below). Palm worm mucus has
been found to contain up to 60% elemental
sulfur on a dry weight basis (Juniper et ai,
1986). The origin of the native sulfur is not
clear. Bacteria appear to be little active in
recently secreted or detrital mucus, both of
which are rich in S° (Juniper, 1988). Detrital
mucus constitutes a significant reservoir of
elemental sulfur at some vents. In one series
of vestimentiferan grabs from Axial Seamount, nearly 25% of the total wet weight
consisted of S°-rich palm worm mucus
(Tunnicliffe et ai, 1985).
177
Influence of Alvinellids on sulfide
mineralization
The abundance of tube building alvinellid
worms on chimney surfaces creates an interesting potential for interaction with mineralization and chimney growth. Near complete coverage of chimney surfaces by
alvinellid tubes has been reported by several
authors (Tunnicliffe and Juniper, 1990;
Juniper et ai, 1992; Chevaldonne and Jollivet, 1993). In this section we will review
what is known of the interaction between
alvinellids and chimney mineralization.
Discussion will be limited to Alvinella spp.
and P. sulfincola.
The influence of Alvinella spp. on chimney mineralization has not been examined
in the same detail as have the effects of P.
sulfincola (see below), although descriptions
of sulfide chimneys do indicate that Alvinella spp. can significantly influence chimney morphology. In their account of the texture of black smoker chimney samples from
East Pacific Rise vents, Haymon and Kastner (1981) note an abundance of interconnected cavities in the outer part of the chimney that are clearly remains ofAlvinella spp.
tubes. The snow ball diffusers of EPR vents
have frequently been referred to as biogenic
edifices created by Alvinella spp. (Fustec et
ai, 1987). No equivalent structures are
known from northeast Pacific vents, even
though the chemistry of the source fluids for
edifice construction is similar (Tunnicliffe
et ai, 1986; Von Damn and Bischoff, 1988;
Butterfield et ai, 1990).
How Alvinella spp. shapes chimney morphology is not known. Local alteration of
mineralogy around Alvinella spp. tubes has
been attributed to interior cooling (Desbruyeres et ai, 1985). As mentioned above,
cooling of the tube may be accomplished by
passive convection if both ends of the tube
are open or by active ventilation by the
worm, as proposed by Chevaldonne et ai
(1991). When worm density is sufficiently
high, cooling of tubes and adjacent chimney
surfaces may be of sufficient magnitude to
influence accretion of chimney material, and
cause some young chimneys to develop into
snow ball structures (Juniper, 1984).
The effect of Paralvinella sulfincola on
178
S. KIM JUNIPER AND PASCALE MARTINEU
FIG. 3. Photo montage showing Paralvinella sulfincola colonies on Fountain chimney (Cleft Segment, Juan de
Fuca Ridge) and removed from its tube ("T" in photo upper right). Worm in insert is approximately 3 cm long.
chimney mineralogy was studied by Paradis
et al. (1988) and Juniper et al. (1992).
Examination of the texture and composition of chimney material beneath worm
tubes consistently revealed a 2 mm-thick
crust of the sulfide mineral marcasite (FeS2)
beneath the mucous tubes (Fig. 5). This suggested that worms altered local geochemistry so as to favor marcasite precipitation.
How might this occur? Formation of both
marcasite and pyrite usually requires the
presence in solution of intermediate sulfur
compounds and FeS precursors; at pH < 5
the precipitation of marcasite is favored over
that of pyrite (Murowchick and Barnes,
1986). High percentage levels of elemental
sulfur have been measured in the tube mucus
of P. sulfincola (Juniper et al., 1986; Juniper, 1988) and similar sulfur concentrations
occur in the tubes of Alvinella pompejana
(Gaill and Hunt, 1986). Paradis et al. (1988)
suggested that interaction of H2S from
hydrothermal fluid with mucus-associated
elemental sulfur could form intermediate
sulfur compounds such as poly sulfide. This
is supported by recent experimental work
on the formation of iron disulfides from
solutions. Reaction of elemental sulfur with
water generates high concentrations of thiosulfate, polysulfides and polythionates at the
sulfur-water interface (Schoonen and Barnes,
1991c). This high local concentration of
intermediate sulfur species apparently can
cause iron disulfides to form directly on elemental sulfur grains (Goldhaber and Stanton, 1987; Schoonen and Barnes, 1991a, b,
c). Thus a local source of elemental sulfur
in tube mucus could cause marcasite precipitation both by direct reaction with FeS
precursors, or indirectly through the generation of intermediate sulfur species.
Marcasite crusts are often continuous over
areas of the chimney surface colonized by
sulfide worms. It has been proposed that the
crusts act as a sealing layer within the outer
chimney wall, reducing inflow of cold sea-
ALVINELLIDS AND SULFIDES
water or outflow of hydrothermal fluid
(Juniper et al, 1992). The sealing effect could
later become important as high-temperature fluids react with the original anhydrite
matrix. The FeS2 layer beneath the worm
tubes, and perhaps the presence of the worm
tubes themselves, would act to reduce
inward draw of seawater through the chimney walls. Because temperature increases
within the chimney conduit cause higher
temperature mineral assemblages to form
(Haymon and Kastner, 1981; Haymon,
1983), this reduced permeability could hasten effects that are normally due to wall
thickening by outwardly growing anhydritesphalerite. The marcasite layer would also
be effective in the opposite direction by
reducing outward seepage of vent fluid. This
in turn would lead to a reduction in temperature and sulfide concentrations in the
worm tubes.
179
obic metabolism have been demonstrated
for several other vent invertebrates
(reviewed in Childress and Fisher, 1992).
Tolerance of anaerobic conditions for periods of several hours appears to be possible
for some vent bivalves and vestimentifera
(Childress and Fisher, 1992). Regular irrigation of alvinellid tubes with sulfide free
seawater could result in exposure to sulfide
being only intermittent, in which case
anaerobic metabolism could be of importance. Turbulent mixing around worm tubes
could also result in only irregular exposure
to sulfide concentrations that would inhibit
aerobic metabolism. With regard to this latter point, both Johnson et al. (1988a) and
Chevaldonne et al. (1991) emphasize the
high variability of temperature for vent
invertebrates and thus sulfide exposure.
External barriers. Recent studies by Powell (1989) and Julian and Arp (1992) indicate that sulfide diffuses across biological
Defenses against sulfide in vent
membranes at rates that are comparable to
alvinellids
oxygen diffusion rates. Alvinellid tubes and
Alvinellid polychaetes live in environ- mucous layers could offer partial protection
ments where ambient mM levels of H2S are against external sulfide by restricting cirat least possible and where levels of 200- culation of vent fluid across epidermal sur300 nM are likely very common (Johnson faces, (see Arp et al. [1995] and Dubilier et
et al., 19886; Butterfield et al., 1990; Sar- al. [ 1995]). Obviously any barrier that would
razin et al., unpublished data). Whether or reduce inward diffusion of H2S would be
not these animals are constantly exposed to similarly impenetrable to oxygen, CO2 and
such levels of sulfide remains to be verified. waste products. One solution to this probAs discussed above, fine scale physico- lem is to be found among non-vent benthic
chemical habitat information for sulfide polychaetes that maintain oxidizing conchimneys is sparse and often unreliable. It ditions within tubes that extend well into
is unlikely that the worms would be able to anoxic sediments, by irrigating their tubes
maintain aerobic metabolism at sulfide con- with seawater (e.g., Aller and Yingst, 1978).
centrations in excess of 300 nM, since aer- There is mineralogical evidence for cooler
obic pathways in all marine invertebrates seawater circulation through tubes of Alvistudied to date are inhibited by such high nella spp., as discussed above, but there is
sulfide concentrations (Eaton and Arp, 1993; no solid evidence to indicate that this occurs
Volkel and Grieshaber, 1994). In addition while the tubes are occupied by living
to diffusion across epidermal surfaces, sul- worms. Concentrations of bacteria on inner
tube walls oiAlvinella spp. (Gaill et al, 1987)
fide can also enter alvinellid tissues across and Paralvinella sulfincola (Juniper, 1994)
the gut wall during digestion of organic mat- are also indicative of distinctive microter associated with sulfide mineral particles. environmental conditions. Whether these
In the following sections, we will summarize conditions are linked to tube irrigation or
what is known of the importance of different to the accumulation of metabolites prosulfide defense mechanisms in alvinellid duced by the worm remains to be deterpolychaetes.
mined. Direct measurement of conditions
Anaerobic metabolism. While anaerobic around and within alvinellid tubes is probmetabolism has not been investigated in the ably required to verify the worms' ability
alvinellids, substantial capacities for anaer-
S. KIM JUNIPER AND PASCALE MARTINEU
180
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B
40
35-
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re
3 Spi
m
2
o
<;
*-»
o>
"5
3025201510-
5-I
0
0
150
300
450
600
Accumulated time (hours)
spire height
anhydrite edge
750
highest worm
ALVINELLIDS AND SULFIDES
181
FIG. 5. Scanning electron micrograph (SEM) of oblique section through uppermost part of Hell Vent chimney
(Axial Seamount, Juan de Fuca Ridge) showing typical mineral profile underlying colonies of Parahinella
sulfincola. From exterior to interior of the chimney (left to right in the photo), there is an outer layer of spongy
marcasite/pyrite with occasional intergrowth of barite needles (BA). Next innermost is a sealing layer of pyrite
(PY) that insulates the outer spongy wall from the hot interior. Remaining material (right side) is a >3 mm
layer of sphalerite (SP), mainly replacing bladed anhydrite. See Juniper et al. (1992) for further mineralogical
detail.
to irrigate and oxygenate their tubes. Mineralogical studies of occupied tubes could
also provide more information about the
tube microenvironment.
The elaborate adaptations of Alvinella spp.
to host epidermal bacteria suggest an
important metabolic role for the epibionts
(Desbruyeres et al., 1983; Desbruyeres and
Laubier, 1986). A nutritional symbiosis was
originally proposed by Alayse-Danet et al.
(1986), but this hypothesis has not yet been
proven. While the potential for sulfide oxidation by these epibionts is unknown, they
could further contribute to maintaining tolerable sulfide levels within the worm's tube
(Alayse-Danet et al., 1987). Apart from very
limited intersegmentary epibiosis of P.
grasslei (Alayse-Danet et al., 1987), epider-
FIG. 4. Time-lapse camera study of Mushroom Vent on Axial Seamount (Juan de Fuca Ridge). A) Small
anhydrite spires growing up from the main body of the sulfide mound were first colonized by Parahinella
sulfincola (upper right) (adapted from Tunnicliffe and Juniper, 1990). B) Summary of data on spire growth,
upward migration of visible sulfide mineralization (anhydrite edge), and vertical migration of sulfide worms
(height of highest worm) (see Juniper et al., 1992).
182
S. KIM JUNIPER AND PASCALE MARTINEU
TABLE 1. Interspecific comparison of sulfide-oxidizing activity between species of Paralvinella and common
intertidal bent hie polychates.*
Sulfide oxidation activity
(imol min- 1 g FW-'
pmo! min" 1 g protein"1
MonobromomobimancHPLC assays
at 200 pM sulfide
(imol min- 1 g protein- 1
0.67 ± 0.53(11)
0.27 ± 0.06 (10)
21.05 ± 12.31(11)
7.65 ± 2.62(10)
1.59 ±0.58(5)
1.33 ±0.62(2)
0.20 ± 0.09 (7)
0.12 ± 0.02 (3)
9.66 ± 4.72(11)
5.36 ± 0.84(3)
1.40 ±0.34(6)
n.d.
Benzyl viologen assays
at 5 mM sulfide
Species
Vent
Paralvinella sulfincola
Paralvinella palmiformis
Non-vent
Nereis virens
Nephtys caeca
Note: Means and standard deviations for a number of samples in parentheses.
* Data in first two columns obtained by following the reduction of an artificial electron acceptor, benzyl
viologen (BV). The method was basically that of Powell and Somero (1985), with whole animal tissue homogenates (without gut contents) being exposed to 5 mM sulfide. Sulfide removal was also verified by the monobromobimane-HPLC method of thiol derivatization (third data column), using an adaptation of the method
Vismann (1991a), where homogenates were exposed to 200 **M H2S. Differences in rates between the two
methods are, in part, a result of different incubation conditions (i.e., sulfide levels, protein concentrations).
mal bacteria have never been described in
any of the known species of Paralvinella. A
sulfide oxidizing capacity also has been suggested for the bacteria that colonize the inner
tube wall of Alvinella spp. (Desbruyeres and
Laubier, 1986, 1991; Hunt, 1992) and P.
sulfincola (Juniper, 1994).
Detoxification in tissues. A capacity to
oxidize sulfide by enzymatic and non-enzymatic (metal ion) catalysis has been identified in a number of benthic invertebrates
(Powell and Arp, 1989; Vismann, 1990,
1991 a, b) and may be quite widespread. This
line of defense would protect haemoglobin
and mitochondria from sulfide diffusing in
from the exterior (Sanders and Childress,
1992; Arp et ai, 1992, 1995). Immobilization of sulfide through binding to metals,
proteins, and glutathione can add to internal
protection (Vismann, 19916; Eaton and Arp,
1993). No data on enzymatic sulfide oxidation or sulfide binding are currently available for Alvinella spp. Some preliminary data
on enzymatic (heat labile) sulfide oxidation
in tissues of Paralvinella sulfincola and P.
palmiformis are presented in Table 1. Interspecific comparisons of sulfide-oxidizing
activity in crude homogenates showed the
higher rate of activity to occur in P. sulfincola (Table 1), although rates for both Paralvinella species were comparable to benthic polychaetes from non-vent habitats.
CONCLUSIONS
The potential influence of vent organisms
on sulfide mineralization is a growing area
of inter-disciplinary research. Alone, observations of textural and mineralogical differences in areas of chimney colonized by
alvinellids, while very suggestive of biological effects, constitute only circumstantial
evidence. There is a clear need for more
supporting biochemical data on sulfide
metabolism, and for information on worm
migration and tube irrigation, before precise
causal mechanisms can be proposed (Juniper et al, 1992; Juniper, 1994). The role of
tube bacteria in the mineralization process
also needs to be further investigated,
especially since there is already a substantial
literature on the influence of vent microorganisms on metal and sulfide oxidizing
reactions (reviewed in Juniper and Tebo,
1995).
Detoxification of sulfide by alvinellids has
received little attention from investigators
in the past, as such studies at vents have
primarily focused on animals with internal
symbionts. The possible role of the Alvinella
spp. epibiotic bacteria in sulfide detoxification still remains to be verified. Critical
to understanding the capacity of vent alvinellids to tolerate or detoxify sulfide are data
on sulfide concentrations that they are actu-
ALVINELLIDS AND SULFIDES
ally exposed to. In this environment of nunscale physico-chemical gradients and turbulent mixing of vent fluids and seawater,
it is presently extremely difficult to describe
the microhabitat of organisms that are only
a few centimeters in length. There is a real
need for instrumentation development that
would enable the same type of microgradient studies to be done at vents as have
been performed in shallow water microbial
mats with micro-electrode technology. The
application of micromanipulators and finescale sensing instruments to the vent environment should lead to a much improved
understanding of how alvinellid worms have
adapted to living in proximity to mM levels
of sulfide and extraordinarily high temperatures.
183
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ACKNOWLEDGMENTS
Pacific: An overview. Biol. Soc. Wash. Bull. no.
This work was supported by NSERC
6:103-116.
Canada through research grants to S. K. Desbruyeres, D. and L. Laubier. 1986. Les AlvinelJuniper. P. Martineu benefitted from postlidae, une famille nouvelle d'annelides polychetes
infeodees aux sources hydrothermales sousgraduate scholarships from FCAR (Quebec)
marines: Systematique, biologie et ecologie. Can
and the GEOTOP Research Centre at the
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Universite du Quebec a Montreal. Graphics Desbruyeres,
D. and L. Laubier. 1991. Systematics,
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phylogeny, ecology, and distribution of the Alvithe organizers of the Life with Sulfide symnellidae (Polychaeta) from deep-sea hydrothermal
posium, Alissa Arp and Chuck Fisher, for
vents. Ophelia Suppl. 5:31-45.
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