Phycologia Volume 53 (5), 426–432 Published 15 September 2014

Chrysophyte stomatocyst production in laboratory culture and descriptions of seven

cyst morphotypes
DALE A. HOLEN
Penn State University, Dunmore, Pennsylvania 18512, USA

ABSTRACT: There is an extensive literature on chrysophyte stomatocysts from diverse habitats and their use as
paleolimnetic indicators of past environmental conditions in lakes. The majority of these cysts are unidentified, and more
laboratory studies are needed to link stomatocyst morphotypes to vegetative stages. A laboratory procedure that resulted
in stomatocyst production in the chrysophyte alga Ochromonas pinguis (Chrysophyceae) was tested to determine if the
protocol would also stimulate the production of stomatocysts in six other chrysophytes. Chrysolepidomonas
dendrolepidota, Chrysosaccus cf. sphaericus, Dermatochrysis sp., Synura cf. americana, Synura cf. macropora and a
second species of Ochromonas all produced stomatocysts within two to three weeks. Morphology of the stomatocyst was
described using scanning electron microscopy (SEM). This was the first report of stomatocysts from Chrysosaccus and the
first SEM description of O. pinguis cysts. These autecological studies provide a greater linkage between sediment
stomatocysts and their biological origin, strengthening the interpretive value of these microfossils and broadening our
understanding of chrysophyte biogeography.
KEY WORDS: Chrysolepidomonas, Chrysophytes, Chrysosaccus, Cyst, Dermatochrysis, Ochromonas, Statospore,
Stomatocysts, Synura

INTRODUCTION
As part of their life history, many algae produce resting cysts
during unfavorable environmental conditions. Although cyst
development is also involved in the sexual cycle of some
algae (Sandgren & Flanagin1986), the production of resting
cysts as a result of environmental stress, including predation
(Rengefors et al. 1998), increases the likelihood of vegetative
cell survival and, in addition, provides a seed bank for
population recruitment when conditions improve. Within the
Chrysophyceae and Synuophyceae, two chrysophyte classes
separated based on biochemical and ultrastructural differ-
ences (Andersen 1987), this resting stage appears particularly
important, as most if not all genera produce silica cysts. The
term stomatocyst is a more specific term for chrysophyte
cysts, but the terms cyst and statospore are also used
interchangeably.
Stomatocysts are typically spherical to ovoid shape, their
outer surface may be smooth or ornamented, and there is a
single pore that is sometimes surrounded by a collar. Viable
stomatocysts have an organic plug that fills the pore. The
significance of silica cysts is exemplified by the high
concentrations of cysts observed in whole lake water
(Firsova et al. 2008) and in sediments (Rybak 1986;
Kamenik et al. 2001; Betts-Piper et al. 2004). Because the
biogeography of many chrysophyte species is influenced by
limnological variables (Sandgren 1988; Siver 1995; Škaloud
et al. 2013), cyst deposition and accumulation in lake
sediments leave a historical record of chrysophyte assem-
blages that can be used as paleolimnological indicators of
past chemical and physical conditions (Adam & Mahood
* Corresponding author (dah13@psu.edu).
DOI: 10.2216/14-001.1
Ó 2014 International Phycological Society
1981; Smol 1988; Kamenik & Schmidt 2005; Pla & Anderson
2005). Nevertheless, only about 5% of chrysophyte stoma-
tocysts have been linked to their vegetative counterparts;
that is, they are unidentified. These unidentified stomatocyst
morphotypes are grouped together and categorized simply as
chrysophyte stomatocysts and assigned numbers as de-
scribed by the guidelines of the International Statospore
Working Group (Cronberg & Sandgren 1986). Although
calibration sets (Charles & Smol 1994) have been developed
for use in models correlating stomatocyst morphotypes to
specific physical and chemical lake conditions (Zeeb & Small
1995; Duff et al. 1997; Wilkinson et al. 1999; Pla et al. 2003),
a clearer picture of chrysophyte distribution patterns would
emerge if vegetative stages and cysts were linked.
In 2008, Ochromonas pinguis Conrad was isolated from an
oligotrophic lake in northeastern Pennsylvania and was
maintained under constant illumination in a temperature-
controlled incubator. When this alga was switched to a 16:8
day/night light cycle as part of a feeding study, stomatocyst
formation was observed. Because there is no prescribed
method for inducing stomatocysts, the intent of this study
was to evaluate the day/night laboratory procedure for
producing stomatocysts in other species. Chrysolepidomonas,
Chrysosaccus, Dermatochrysis, two Synura taxa and a second
species of Ochromonas were subjected to the same laboratory
regiment as O. pinguis. Stomatocysts were produced in all
cultures. This article describes the specific laboratory
technique and scanning electron microscopy (SEM) of the
stomatocysts.
MATERIAL AND METHODS
Ochromonas pinguis strain Giles 1 was isolated from Lake
Giles, an oligotrophic lake located in northeastern Pennsyl-

426

vania, and Ochromonas sp. strain Sem 1 was isolated from
Sweeny Pond, a small brown-water reservoir (0.052 km2)
also located in northeastern Pennsylvania (Williamson et al.
1997; Holen 2010). Chrysolepidomonas dendrolepidota Peters
et Andersen strain CCMP 293, Dermatochrysis sp. strain
A12,609, Chrysosaccus cf. sphaericus Bourrelly strain CCMP
1162 Synura cf. americana Kynčlová & Šlakoud strain
A12,391 and Synura cf. macrospora Šlakoud & Kynčlová
strain A12,220 were kindly provided by Dr. Robert
Andersen. All cultures except Synura cf. americana, which
died, are available from the author upon request.
Prior to the experiment, nonaxenic cultures were main-
tained on DY-V medium (Andersen et al. 2005) at 258C on a
16:8 day/night light cycle at 100 lmol m2 s1. To induce
stomatocyst formation, a 1-ml sample in stationary phase of
growth was added to 50 ml of sterile-filtered DYV medium
in 125-ml glass, sterile Erlenmeyer flasks. The foil-topped
flasks were placed in a temperature-controlled incubator set
at 218C on a 16:8 day/night cycle and illuminated from one
side by a bank of cool-white fluorescent bulbs at 100 lmol
m2 s1. The cultures were manually shaken daily. Cyst
production was monitored using visual observation every
two to three days by scanning a wet mount at 3600 and
31000 using an Olympus BX51 microscope fitted with 360
UPlanFLW and 3100 UPlanApo objective lenses (Olympus
Corp., Tokyo, Japan). If no stomatocysts were observed in
wet mount preparations, then a 5-ml subsample was pelleted
by centrifugation at 1800 3 g. The supernatant was discarded
and the pellet vortexed in 30% H2O2 and immersed in a 508C
water bath for 24 hours following the procedure of Battarbee
(1986). Samples were sedimented by centrifugation, the pellet
was resuspended in distilled water, and a small aliquot was
removed to scan microscopically for stomatocysts at 3600 to
31000. Once stomatocysts were observed in a culture, a 10-
ml subsample was removed and treated with H2O2 as
described above for SEM preparation.
For SEM, the samples were collected on a 0.22-lm-pore-
size 13-mm-diameter Millipore polycarbonate filter (Merck-
Millipore Corp., Billerica, Massachusetts USA). The filter
was transferred to 2.5% glutaraldehyde in 0.1 M sodium
cacodylate buffer for 1 h at room temperature. The filter was
washed three times for 5 min each using 0.1 M sodium
cacodylate buffer; the filter was then dehydrated in a graded
ethanol series (50%, 70%, 85%, 90%, 95%, 33 100%) for 5
minutes each. Samples were critical point dried, mounted on
studs and sputter coated with 10 nm Au/PD. Samples were
examined using a JEOL JSM 5400 scanning electron
microscope (JEOL Ltd, Tokyo, Japan) operated at 20 kV.
RESULTS
The mature stomatocyst of O. pinguis was spherical to slightly
oval with a length and width of 17.4–19.2 lm and 15.7–16.9
lm, respectively (n ¼ 21; Figs 1, 2). The wide secondary collar
was cylindrical and complex with a mean height of 2.48 6 0.28
lm (n¼10), and a thick, concave annulus separates the primary
and secondary collars. The mean collar width is 8.5 6 0.36 lm
(n ¼ 15) and is reflexed at its base. The 1-lm pore appeared to
be regular and was often observed with a plug in mature
Holen: Inducing chrysophyte stomatocyst formation 427
stomatocysts. The stomatocyst’s surface was heavily orna-
mented with thick lunate ridges of varying lengths and heights.
These projections ranged from short verrucae to long, curvy
ridges. Some of the ridges were bifurcated and others so
prominently elevated as to appear more plate-like in appear-
ance. The ridges were uniformly distributed over the surface of
the stomatocyst and are slightly thicker at their base. Assuming
a random distribution over its surface, the mean number of
these projections on the stomatocyst was estimated to be 57 6
6.1 (n ¼ 12).
Ochromonas sp. was an undescribed species. The vegeta-
tive cell was small (4–6 lm diameter) and pleomorphic; the
typical cell shape was spherical to oval with a wide anterior
flagellar depression that gave it a heart-shaped appearance.
The anterior side, over which the longer flagellum arcs, was
more rounded and slightly elevated. The long and short
flagella were approximately 8 lm and 2 lm in length,
respectively. The chloroplast was ribbon-like and positioned
off center from the more rounded anterior side of the cell to
the posterior end of the cell; the plastid was sometimes
twisted. There was a noticeable stigma. The shape of the
stomatocyst was spherical to oval (5.6–9.6 lm long, 3.8–8.3
lm wide) with a distinct collar (n ¼ 10; Fig. 3). The mature
cyst was smooth (devoid of ornamentation) and had a simple
conical collar that arose abruptly from the cyst body. The
conical collar had a basal diameter of 2.3–2.9 lm with an
apex diameter of 1.3–1.5 lm and a collar height of 0.8–1.2
lm (n ¼ 10). A plug was never observed. The collar had a
rounded apex and a regular pore with a pore diameter of
0.2–0.4 lm (n ¼ 10). Collar morphology was variable; some
cysts exhibited a true collar that entirely encircled the pore,
but other cysts possessed a collar that was interrupted and
left a small break or groove that was continuous from its
apex to the base (Fig. 4).
Dermatochrysis sp. had a spherical stomatocyst with a
smooth surface during its early stage of development, but the
cyst surface was uniformly scabrous (approximately 0.1 lm
in height) on mature cysts (Figs 5, 6). The average cyst
diameter was 6.6 lm and ranged from 5.9 lm to 7.2 lm (n ¼
5). There was no collar development. The cyst contained a
regular conical pore with an inner and outer diameter of 0.4–
0.5 lm and 0.8–1.0 lm, respectively (n ¼ 5).
The cyst of C. dendrolepidota was spherical to slightly
oblate and had a smooth surface. The cyst diameter was 6.1–
6.8 lm. There was a regular pore that appeared flush with
the cyst body (n ¼ 3; Fig. 7). The pore appeared unusually
small in comparison to the cyst body and ranged from 0.4
lm to 0.5 lm in diameter (n ¼ 3). The slight anterior
depression in the oblate-shaped cysts gave the impression of
a narrow lip surrounding the pore opening (Fig. 8).
Synura cf. americana produced a stomatocyst that was
usually spherical with a diameter of 8.4–8.9 lm (n ¼ 3), but
oval cysts have been observed (Fig. 9). The cyst surface was
unornamented and had a shallow concave pore with an inner
and outer diameter 0.5 lm and 1.5–2.0 lm, respectively. The
identity of the strain was determined using SEM observa-
tions of silica scales (Figs S1, S2; Table S1). Unfortunately,
the culture died, and further study was not possible.
Synura cf. macropora had spherical stomatocysts that were
8.8–9.8 lm diameter (n ¼ 9). Cysts had a smooth surface and
a shallow concave pore (Fig. 10). The depressed pore had an
428 Phycologia, Vol. 53 (5)

Fig. 1. Stomatocyst of Ochromonas pinguis strain Giles 1. Note regular pore (arrow). Scale bar ¼ 10 lm.
Fig. 2. Stomatocyst of Ochromonas pinguis. Note reflex at the base of the secondary collar. Scale bar ¼ 10 lm.
Fig. 3. Stomatocyst of Ochromonas sp. strain Sem 1 with complete collar. Scale bar ¼ 2 lm.
Fig. 4. Stomatocyst of Ochromonas sp. with incomplete collar. Scale bar ¼ 2 lm.

inner diameter of 0.6–0.8 lm and an outer diameter of 1.8–
2.0 lm (n ¼ 7). Identity of the strain was based on SEM
observations of the silica scales (Figs S3, S4; Table S1);
however, the scales had significantly smaller keel pores.
Chrysosaccus cf. sphaericus produced a smooth stomato-
cyst that was generally spherical with a diameter of 4.5–6.7
lm (n ¼ 8); however, some cysts appeared slightly ovoid in
shape (Fig. 11). The cyst had a pronounced conical collar,
and, like Ochromonas sp., some cysts were observed with
incomplete collars (Fig. 12). There was an abrupt delineation
between the cyst body and the collar. Complete conical
collars had an acute apex with a sloping inner margin that
surrounded the regular pore. Incomplete collars exhibited a
more rounded apex that encircled approximately three-
fourths of the pore with each collar margin sloping
asymmetrically toward the cyst body. Collar height ranged
from 0.5 lm to 0.7 lm (n ¼ 4). The collar base and apex
diameters ranged from 2.0 lm to 2.4 lm and 1.1 lm to 1.4
lm, respectively (n ¼ 4), and the pore diameter ranged from
0.3 lm to 0.8 lm (n ¼ 3).
DISCUSSION
All seven chrysophytes produced stomatocysts, but it is not
clear which induction stimuli were responsible for their
formation. Removing 5 ml for centrifugation from the
Ochromonas cultures concentrated the stomatocysts and
expedited their detection. Preliminary work showed that the
encystment frequency (percentage of cells producing stoma-
tocysts) for O. pinguis was extremely low, and a similar low
cyst percentage was found here for Ochromonas sp. Curtin
(1987) observed that, on average, 0.006% of the population
of Ochromonas minuscula Conrad encysted regardless of
culture conditions, although the percentage increased
slightly during its stationary phase of growth.
Doflein (1923) was the first to describe the stomatocyst of
Ochromonas pinguis, which at that date he named O. crenata
Klebs. Because of the limits of light microscopy, the cyst
surface was described as being ornamented with spines. The
present study provides the first SEM images of the O. pinguis
stomatocyst, and the apparent spines observed in light
microscopy are shown to be lunate ridges that are slightly
thicker at the base. The stomatocysts of O. pinguis resemble
cyst 133 (Duff et al. 1995), but O. pinguis cysts are larger.
The O. pinguis cysts are also nearly identical to the cyst
produced by Ochromonas sphaerocystis Matvienko (Ander-
sen 1982), but there are differences in collar height (’ 0.6 lm
in O. sphaerocystis vs 2.5 lm in O. pinguis). Because cyst 133
resembles cysts of both O. pinguis and O. sphaerocystis, cyst
133 may be produced by Ochromonas or an Ochromonas-like
organism.
Morphologically, the small Ochromonas sp. flagellate
resembles Ochromonas minima Throndson. However, Flöder
et al. (2006) reported that O. minima was not to be able to
Fig. 5. Immature stomatocyst of Dermatochrysis sp. strain A12,609 with unornamented surface. Scale bar ¼ 1 lm.
Fig. 6. Developmentally mature stomatocyst of Dermatochrysis sp. with scabrae. Scale bar ¼ 1 lm.
Holen: Inducing chrysophyte stomatocyst formation 429
grow in the absence of light; whereas, this flagellate exhibits
a higher growth rate in the dark than the light when fed
bacteria (Holen 2010). It also resembles O. minuta (I.F.
Lewis) Bourrelly except for the length of the small flagellum,
which in O. minuta is exceptionally short in comparison to
the longer flagellum (Pringsheim & Pringsheim 1959). The
shape and location of the chloroplast of Ochromonas sp. is
different from that for Ochromonas minuscula, which has a
more cup-shaped plastid that is located in the cell anterior
(Starmach 1985; Curtin 1987). Moreover, the stomatocyst of
O. minuscula has been previously described, and, although
similar in cyst-body morphology, the collar shape, height
and width are significantly different (Curtin 1987).
Stomatocyst 67 (Duff et al. 1995) is most certainly the
same as described here for Dermatochrysis. Both stomato-
cysts have scabrae uniformly distributed over their surface
and lack a collar. The diameter of Dermatochrysis cysts (5.9–
7.2 lm) is equivalent to stomatocyst 67 with a diameter of
7.9 lm, and pore diameters are similar. Phaeosphaera
perforata Whitford (Whitford 1946) was later considered to
be identical to Tetrasporopsis reticulata Meyer (Whitford
1976), and T. reticulata is the basonym for Dermatochrysis
reticulata (Meyer) Entwisle & Andersen. Whitford (1946)
describes the stomatocyst as smooth. Either the scabrae were
unobserved by Whitford, or his description was based on an
immature form like Fig. 5; alternatively, my observations on
Dermatochrysis sp. may indicate variation within the genus.
Cysts were not previously reported from Chrysosaccus,
and this study augments our knowledge of the genus by
describing its stomatocyst formation. Chrysosaccus cf.
sphaericus was previously named Chrysosaccus cf. epilithicus
Starmach (Andersen 2007).
My description of C. dendrolepidota stomatocysts concurs
with the description reported by Peters & Andersen (1993).
The stomatocysts for the two Synura species in this study
were indistinguishable from each other; however, they were
considerably smaller than those described by Sandgren &
Flanagin (1986; 8–9 lm vs 13–16 lm). Minor stomatocyst
size variation is natural; for example, differences in the
nutritional state of the vegetative cell populations and/or
concurrent cell size at the time of stomatocyst formation may
explain size variation, but this area is not extensively studied.
Similar intraspecific size variations in stomatocysts have also
been reported by Cronberg (1988) and Findenig et al. (2010);
however, in this case, stomatocyst difference could also
reflect differences between the two S. petersenii-like cryptic
species (Škaloud et al. 2012).
Not all stomatocysts are easily identifiable because many
chrysophytes produce simple unornamented cysts that lack a
collar and are morphologically indistinguishable. For
example, the stomatocysts of Synura (Figs 9, 10) are
morphologically similar, and they also resemble the imma-
ture cysts of Dermatochrysis (Fig. 5) and Chrysosphaerella
longspina Lauterborn (Sandgren 1989). Moreover, the
collared and unornamented cysts of Ochromonas sp. and
430 Phycologia, Vol. 53 (5)

Fig. 7. Stomatocyst of Chrysolepidomonas dendrolepidota strain CCMP 293. Scale bar ¼ 2 lm.
Fig. 8. Stomatocyst of Chrysolepidomonas dendrolepidota. Note the anterior depression encircling the pore giving the impression of a small
conical collar or lip. Scale bar ¼ 2lm.
Fig. 9. Stomatocyst of Synura cf. americana strain A12,391. Scale bar ¼ 2 lm.
Fig. 10. Stomatocyst of Synura cf. macropora. strain A12,220. Scale bar ¼ 2 lm.
Fig. 11. Stomatocyst of Chrysosaccus cf. sphaericus strain CCMP 1162 with complete collar. Scale bar ¼ 2 lm.
Fig. 12. Stomatocyst of Chrysosaccus cf. sphaericus with incomplete collar. Scale bar ¼ 2 lm.

Chrysosaccus are also indistinguishable from each other; in
size and appearance, both resemble stomatocyst 126 (Duff et
al. 1995). Stomatocyst 190 (Wilkinson et al. 2001) is also
morphologically similar except in size (8.4–13.5 lm in
diameter). Size could be a useful criterion for differentiating
between developmentally similar stomatocysts but would
require extensive laboratory-based studies of individual taxa
to determine the within-species natural size variation under
different environmental conditions.
The morphology of the pore-collar complex in chryso-
phytes can vary extensively, but because its shape is believed
to be conservative within a species, it is regarded as an
important taxonomic marker for cyst identification (Sandg-
ren & Carney 1983). In both Ochromonas sp. and
Chrysosaccus, the cyst-collar complex varied from being a
true collar that completely encircles the pore to one that is
incomplete with breaks of varying size. Findenig et al.
(2010) also noted that the collars of the colorless chryso-
phyte Spumella vulgaris Cienkowsky (strain 199hm) varied
extensively in laboratory-cultured individuals with some
collars exhibiting short interruptions. The incomplete collar
form may simply represent stomatocysts in which collar
development was interrupted at the time samples were
collected and fixed. However, this phenomenon is not
limited to laboratory samples, as Siver (1991) observed
Mallomonas acaroides v. muskokana Nichols cysts exhibiting
varying degrees of morphological maturity in a core sample
from Little Echo Pond in New York, USA. Whether these
variant cyst morphologies represent different stages of cyst
development or are actually indicative of a natural variation
that exists in cyst development, as has been reported by
Sandgren (1983), needs further investigation. This type of
variation certainly speaks to the difficulty of cyst identifi-
cation in field samples.
Clearly, further autecological studies on known chryso-
phytes are necessary not only to reduce taxonomic uncer-
tainties but also to delineate the environmental cues
pertinent to stomatocyst induction. Coupling these data
with the physical and chemical optima of stomatocysts
morphotypes will provide a broader understanding of the
environmental tolerances and distributions of individual
chrysophyte species.
ACKNOWLEDGEMENTS
I would like to acknowledge and thank Missy Hazen and
John Cantolina at The Huck Institute of the Life Sciences
Electron Microscopy Facility for preparing the stomatocysts
and scales for SEM. I would also like to thank Dr. Robert
Andersen for supplying some of the chrysophytes in this
study. Lastly I thank the reviewers of this manuscript for
their valuable input.
SUPPLEMENTARY DATA
Supplementary data associated with this article can be found
online at http://dx.doi.org/10.2216/14-001.1.s1.
Holen: Inducing chrysophyte stomatocyst formation 431
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Received 2 January 2014; accepted 20 May 2014
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