Development of the Sexual Phase of Pseudocolysis bradeorum (Polypodiaceae)

13 Pages·2001·6 MB·English

by  Blanca Pérez-García

American
Fern Journal 91(4]:214-226 (2001)
Development
of the Sexual Phase of Pseudocolysis
bradeorum
(Polypodiaceae)
Blanca Perez-Garcia, Aniceto Mendoza, Ramon Riba
Autonoma
Universidad Metropolitana, Iztapalapa. Depto de Biologia
A.P. 55-535 C. 09340. Mexico, D.
P. F.
Gomez-Pignataro
Luis D.
Organization for Tropical Studies, OTS, Costa Rica
—
Abstract.^ The development and morphology of the sexual phase of Pseudocolysis bradeorum
(Polypodiaceae) are described from material collected in Finca La Selva, near Puerto Viejo, Prov-
ince of Heredia, Costa Rica. Spores were sown in Thompson medium with agar (25 Petri dishes)
and germinated after seven days; the germination pattern was Gleichenia-iype and the prothallial
development
Dry^naria-lype. Gametangia were typical of homosporous leptosporangiate Spo-
ferns.
rophytes appeared after seven months of culture. The sexual phase of species shares many
this
New
morphological characteristics with Old and World species of Polypodiaceae. There a ten-
is
dency
for vegetative propagation with Pseudocolysis gametophytes.
Pseudocolysis bradeorum Gomez
(Rosenst.) L. D. (Polypodiaceae) a rare
is
growing
terrestrial fern in tropical rain forest soils rich in organic matter, main-
Q
Mexico
(Veracruz, Chiapas), Belize, Nicaragua, Costa and Panama. have
Rica, Plants
long-creeping rhizomes and sub-dimorphic dimorphic
to leaves. Vegetative
Sim
pinnatifid with long petioles. Venation anastomosing with included
veinlets
is
within the
areoles. Sori are regularly to irregularly linear, oblique and exin-
are
some =
Colysis spp. Leptochilus, Flora Malesiana, and
1998] Pleopeltis
vol.
3,
[
and
(Evans Mickel, Gomez,
1969;
1977).
Tryon and Tryon
(1982) placed Pseudocolysis in the subtribe Polypodieae,
but with uncertain Moran &
relationships. Riba suggested probable
(1995) a
hybrid because
origin of the highly variable lamina, the variable petiole length,
and
the occasionally interrupted
sori.
Development
of the sexual phase bradeorum unknown,
of although
P. is
there information about morphology
is the of the gametophytes from other
genera of Polypodiaceae. For example, Nayar
(1962) studied several species of
Polypodium-
the group
Pleopeltis [Arthromeris tenuicaudata (Hook.) Ching,
A. wallichiana (Spreng.) Ching, Campyloneurum
angustifolium Co-
(Sw.) Fee,
lysis elliptica (Thunb.) Ching, hemionitidea pedunculata
C.
(Wall.) C. Presl, C.
(Hook, Grev.) Ching, Crypsinus
et griffithianus (Hook.) Copel., Crypsinus has-
Lemmaphyllum
tatus (Thunb.) Copel., carnosum
(Hook.)
C. Presl, Pleopeltis
excavata (Bory ex Willd.) T. Moore, Pleopeltis normalis Moore, and Poly-
T.
amoenum
Dodium
Wall.l.
UM
BRADEOR
215
Prothalli of about 20 genera of Polypodiaceae have been described by
several
authors (Nayar 1954, ig63a, 1963b, 1964, 1965; Bajpai 1964; Nayar and Chan-
dra 1965; Nayar and Kaur 1969, 1971; Schmelzeisen 1933]. Authors reported
that, in Polypodiaceae, the development and morphology
of the prothalli are
perhaps more
understanding
significant for the of evolutionary processes, in
the polypodiaceous ferns than for other fern groups.
Nayar and Raza studied development and morphology
(1970] prothallial of
three species of Pleopeltidoideae [Lepisorus loriformis (Wall.] Ching, thun-
L.
bergianus Ching and Weatherbia accedens (Blume] and one
(Kaulf.] Copel.],
species of Polypodioideae [Polypodium However, mor-
vulgareL.]]. prothallial
phology most and
for species of Pleopeltidoideae Polypodioideae poorly
is
known. Most which
taxa for the prothallus has been described belong to sub-
and
families Microsorioideae, Crypsinioideae, Platycerioideae.
New
Several studies are important for our understanding of gametophytes of
World polypods, ones by Stokey on Polypodium
in particular (1945, 1954]
R
pectinatum and plumula Humb. & Bonpl. ex Atkinson & Stokey
L. Willd.;
(1964, 1970] on Polypodium chnoodes Spreng.; Reyes and Perez-Garcfa (1994]
on Polypodium lepidotrichum Maxon; Reyes on Niphidium
(Fee] (1996]
et al.
on
crassifolium Lellinger; Perez-Garcia (1998] three species of
(L.] et al. Pbile-
bodium Brown] Smith; and Ramirez and Perez-Garcia on Micro-
(1998]
(R.
J.
gramma
nitida Sm.] A. R. Sm. (see comparative Table
1].
(J.
An known
attempt to classify the various patterns of spore germination in
Momose
homosporous ferns was made by (1942], he described three types of
spore germination: centrifugal, centripetal, and tangential. Later, Nishida
(1965] renamed the centripetal and tangential types aspidiod and polypodiod,
respectively, and added another which he called the tripolar type. Recently,
Nayar and Kaur (1968] pointed out that these interpretations of spore germi-
nation are incorrect inasmuch as they ignored the polarity of the germinating
spore. Nayar and Kaur (1971] gave a detailed account of the patterns of spore
germination, classifying them on the basis of the planes of cell division in
and growth primary
relation the polarity of the spore directions of of the
to
Thus among homosporous
rhizoid and prothallus. ferns there are three distinct
and
equatorial and amorphous, differing in the plane
categories: polar, all
sequence
of divisions.
cell
The prothallus of the homosporous ferns follows a definite pattern of de-
velopment leading ultimately to the characteristic adult form. This pattern is
commonly
constant each species and to taxa of higher order. In general
for
among
development recognized ho-
seven different patterns of prothallial are
mosporous by Nayar and Kaur (1969]:^dia/7fum-type, Ceratopteris-type,
ferns
and Osmunda-iype. These differ in the sequence
Drynaria-type, Kaulinia-type,
which
development and the region a mer-
of cell divisions, in the stage of at
and form
istem presence or absence of trichomes, in the final
established,
is
of the adult
thallus.
which
Thus the type of prothallial development in an apical cell is estab-
[Adiontum-type] ap-
lished early during the formation of a prothallial plate
pears be primitive compared to those in which the establishment of a :r.er-
to
216 FERN
Table
Comparison
1. of the different development stages of the gametophytes of the some genera of
the Polypodiaceae family.
Types
of spore
Prothallial
Tax a Type of spore germination Filamentous phase development
Pseiidocolysis tra- Monolete,
yellow. Gleichenia-type Germinal
filaments Drynaria-type
^r ^v ^^_
deorum
ovate to elliptic of 2-3
cells
long
No
^Arthromeris tenui- Monolete, brown, mentioned
Germinal Young
filaments
prothalli
Cauda
piano convex or of 3-7 more
cells spatulate
concavo convex
long
or less strap-
shaped
a cordate
J
^Arthromeris No
walli- Monolete, dark mentioned Germinal Young
filaments
prothalli
chiana brow
piano 3-7
n, of more
cells spatulate
convex
or con-
long
or less strap-
cavo convex
shaped a cordate
Campykmeurum
No
' Monolete, golden mentioned Germinal filaments Young
prothalli
^
I.
angusiifolium yellow, piano 3-7
of
cells spatulate
convex
or con-
long
cavo convex
^Colysis eUiptica Monolete, golden No mentioned Germinal Young
filaments
prothalli
yellow, promi- 3-7
of more
cells spatulate
nently concavo-
long
or less strap-
convex
shaped
a cordate
No
^Colysis hemioniti' Monoloete, golden
mentioned Germinal Young
filaments
prothalli
dea
yellow
promi-
3-7
of mote
cells spatulate
nently concavo-
long
or less strap-
convex
shaped
a cordate
^^^^^_ ^_ ^^ ^^ ^^
^ri Colysis peduncula- Monolete, golden No mentioned
Germinal Young
filaments
prothalli
ta yellow, promi-
3-7
of more
cells spatulate
nently concavo-
long
or
less strap-
convex
shaped
a cordate
^Crypsinus No
Monolete,
gnffithi- pale mentioned Germinal Young
filaments
prothalli
ajius brown, promi-
8-
of 10
cells Strap-shaped a
nently concavo-
^
long
cordate
convex
^Crypsinus hastatus Monolete, No
pale mentioned
Germinal Young
filaments
prothalli
brown,
promi-
8-10
of
cells strap-shaped a
nently concavo-
long
cordate
convex
^Lemmaphyllum
Monolete, brown- No
mentioned
Germinal Young
filaments
prothalli
carnosum
ish yellow,
pla- 3-7
of
cells spatulate
no-convex
or
long
slightly conca-
vo-convex
^Pleopeltis excavata Monolete, No
golden mentioned
Germinal Young
filaments
prothalli
yellow; plano-
of 3-7
cells cordate
convex
or
long
^p^
slightly conca-
vo-convex
norma
Pleopeltis Monolete, No
^ lis golden mentioned Germinal Young
filaments
prothalli
yellow,
plano-
of 3-7
cells cordate
convex
or
long
slightly conca-
vo-convex
BRADEORUM
PEREZ-GARCIA ET PSEUDOCOLYSIS SEXUAL PHASE
AL.: 217
Table
Extended.
I.
Prothallial hairs Adult prothallus Sexual expression Sporophytes
No
hairs Cordiform-spatulate 6 and 9 dioic, 9 7 months
,
4-
^ * type usual, neck
6 long
cells
No
Unicellular papillate with Cordate with midrib small, 9 type usu- sporophyte
(?
4-8 4-6
neck
a extracellular cap-like cells thick cells
al,
secretion long
No
6
Unicellular papillate with Cordate with midrib small, 9 type usu- sporophyte
4-6
4 neck
a extracellular can-like 8 cells thick cells
al,
long
secretion
No
Unicellular papillate with Cordate w^ith midrib S small, 9 type usu- sporophyte
4-6
4-8 neck
a extracellular cap-like cells thick al, cells
long
secretion
No 6 No
hairs Perennial, ribbon- small, 9 type usu- sporophyte
4-6
shaped branched neck
al, cells
long
No
No Perennial, ribbon- S small, 9 type usu- sporophyte
hairs
4-6
shaped branched neck cells
al,
long
No
No hairs Perennial, ribbon- S small, 9 type usu- sporophyte
4-6
shaped branched neck cells
al,
long
No
S sporophyte
Cordate with midrib small, 9 type usu-
Unicellular papillate v\ith
5-6
4-8 neck
a extracellular cap-like cells thick al, cells
long
secretion
No
Unicellular papillate with Cordate with midrib S small. 9 type usu- sporophyte
5-6
a extracellular cap-like 4-8 cells thick al, neck cells
long
secretion
No
sporophyte
Unicellular papillate with Cordate with midrib 3 globular, 9 type
4-6
4-8 neck
thick usual,
a extracellular cap-like cells
long
cells
secretion
No
sporoph)te
Unicellular papillate with Cordate with midrib 6 small, 9 type usu-
4-6
4-8 neck cells
a extracellular cap-like cells thick al,
long
secretion
No
sporophyte
Branched more com- Ribbon-shaped S small, 9 type usu-
or
4-6
neck
cells
plex al,
long
218 ERN
Table
Continued.
I.
Types of spore
Prothallial
Taxa Type of spore germination Filamentous phase development
Polypodium am- No
Monolete, hyaline mentioned Germinal Young
filaments
prothalli
moeniun
to yellowish, 8-10
of
cells spatulate
plano-convex
or long
slightly conca-
vo-convex
^Pseudodrynaria Monolete No
hyaline mentioned Germinal Young
filaments
prothalli
coron cms
3-4
of
cells spatulate
long
LTepisorus
•^ ^^?^i c lorifor- Monolete, yellow- Vittaria-type Germinal
filaments Kaulinia-type,
mis
ish globules ^-5
oil of young
cells prothalli
long
elongate
a rib-
bon-like
^Lepisorus thimber- Monolete,
yellow-
Vittaria-type Germinal Dry
filaments n aria- type,
giamis
ish 4-5
oil ^ilobules of young
cells prothalli
long
elongate
a near-
ly strap-like
^Weatherbya
acce- Monolete,
yellow-
Vittaria-type Germinal
filaments Drynaria
type.
dens
ish oil globules of 6-8 young
cells prothalli
long
cordate
^Polypodium
vidga- Monolete, yellow-
Vittaria-type Germinal
filaments Drynaria-type,
re
ish globules 6-8
oil of young
cells prothalli
long
cordate
^Polypodium pectin- Monolete No
mentioned No
Germinal filaments mentioned
citum
4-8
of
long
cells
"^Polypodium plumu- Monolete No
mentioned No
Germinal
filaments mentioned
la
4-8
of
Ions
cells
Polypodium chnoo- Monolete, brown No
mentioned No
Germinal
filaments mentioned,
des
3-4
of
long
cells spatulate plate
more common,
3-6
long
cells
^Polypodium lepido- Monolete, brown No
mentioned No
Germinal
filaments mentioned.
trichum
4-5
of long
cells spatulate plate
'^Niphidium crassi- Monolete, yellow-
Vittaria-type Germinal
filaments Drynaria-type,
folium green
4-7
of
long
cells spatulate plate
^Phlebodium
ara- Monolete, golden
Vittaria-type Germinal
filaments Drynaria-type,
neosum
2-7
of
cells spatulate plate
+
long
^Phlebodium
decu- Monolete, golden
Vittaria-type Germinal
filaments Drynaria-type,
mamim
of 2-7
cells spatulate plate
long
^Phlebodium pseu- Monolete, golden
Vittaria-type Germinal
filaments Drynaria-type,
doaureum
of 2-7
cells spatulate plate
long
"^Micrognunma
niti- Monolete, yellow
Vittaria-type Germinal
filaments Drynaria-type,
da
4-6
of
long
cells spatulate plate
Nayan &
1962; Nayar, 1954; ^Nayar Raza, &
' ^ 1970; ' Stokey, 1959; ^ Atkinson Slokey, 1970; ^ Reyes
&
Perez-Garcia, 1994; ' Reyes et aL, 1996; « Perez-Garcia et aL, 1998; ** Ramfrez & Perez-Garcfa, 1998.
= =
S
anteridia, 9 archesonia.
PEREZ-GARCIA ET PSEUDOCOLYSIS BRADEORUM SEXUAL PHASE
AL.: 219
Table
Continued. Extended.
1
.
Prothallial hairs Adult prothallus Sexual expression Sporophytes
Unicellular papillate with Cordate with midrib 6 cap cell large, basal No sporophyte
4-8
a extracellular cap-like cells thick cell columnar, bar-
secretion rel-shaped, rather
than saucer or fun-
nel-shaped
No
Unicellular papillate Cordate S globose and sessile, sporophyte
2 months 9 type
4-5
neck
usual, cells
3-4 month
long,
No
Unicellular papillate se- Elongate, ribbon-like T^pe usual in Polypo- sporophyte
cretory diaceae, neck short
3-4
cells
No
Unicellular papillate se- Elongate cordate Type usual in Polypo- sporophyte
neck
cretory diaceae, short
3-4
cells
No
Unicellular papillate se- Cordate Type usual in Polypo- sporophyte
neck
cretory diaceae, short
3-4
cells
No
Unicellular papillate se- Cordate Type usual in Polypo- sporophyte
neck
cretory diaceae, short
3-4
cells
Cordate S and 9 type usual, Sexual
Unicellular
2-4
months
No S and 4 months, embryo apogamous
Cordate 2
hairs, only hairs
months
gamcto- type usual 3
acicular
in
apogamic
phytes
Cordate 6 globose and elon- Sexual
Unicellular papillate in
3-4
neck
margin, branched gated, $
hairs
long
surface cells
in
3-4
Unicellular simple Cordiform and reni- S globose 3 cells and Sexual, months
form 2 usual, dioic
Cordiform with wide S globose, 9 necks 4 Sexual, 3 months
Unicellular papillate
wings long
cells
small, spherical or Sexual
Unicellular capitate Cordate-spatulate to (?
subglobose, 2
cordate-rcniform
necks 5-7 long
cells
S small, spherical or Sexual
Unicellular capitate Cordate-spatulate to
subglobose, 9
cordate-reniform
5-7
necks long
cells
S small, spherical or Sexual
Cordate-spatulate to
Unicellular capitate
subglobose, 9
cordate-reniform
5-7
necks long
cells
Sexual
No Monoic, small bar-
Cordiform-alargate
hairs cJ
9 necks 5
rii-shape,
lone
cells
220 AMERICAN FERN TOURNAL: VOLUME
istematic cell is delayed [Drynaria-iype]. Usually accompanying delayed
this
establishment
of the meristematic
cell there is also a distinct reduction in
its
an
ma
Dry.
matic cell does not play a very a „_
^.^ ^^ ....^^..^.w.. „. ...^
young
prothallus, as is the case in the majority of the Polypodiaceae (Nayar,
1962, 1963a,
1965).
The
elimination of the meristematic and
cell altogether the establishment of
a pluricellular meristem from some
directly of the marginal cells of a broad,
non-meristic
prothallial plate, has been recorded in several cases among the
more advanced
groups homosporous The
of
ferns. elimination of meristic
cells
and
the presence of a pluricellular meristem found
as in the Kaulinia-iype of
development
appears represent most advanced
to the condition.
we
In this paper, describe the morphology and development
of the sexual
phase
of Pseudocolysis bradeorum.
Materials and Methods
Spores bradeorum
of were obtained from specimens
P. fertile of plants [Go-
mez
26045, USJ) growing Finca La
in Selva, near Puerto province
Viejo, of
Heredia, Costa an
Rica, at elevation of 50 m. Fertile leaves were sealed in paper
envelopes and allowed dry under
to natural conditions
to favor the release of
spores from
the sporangia. After several days, the contents of the envelopes
were
sifted to eliminate sporangial fragments and
other debris (Perez-Garcfa
et 1998).
al.,
Spores were sown
in 20 Petri dishes (two on 10%
replicates) agar supple-
mented Thompson medium
with
(Klekowski,
1969; Perez-Garcia, one
1989);
which was
of
kept darkness
in to test for photoblastism. Petri dishes were kept
in transparent, sealed plastic bags to avoid dessication and contamination,
given a regime h h
light of 12 light/12 darkness, with lamps
daylight Solar
cm
{75w/t 38/Al-D) placed
35 above the cultures, and maintained temper-
at
ature 25-28°C. Cultures were examined
periodically (except the dish
Petri
maintained
in darkness) until germination was on
detected seventh
the day.
After that, dishes were checked every 15 days to record data on the develop-
ment
of the prothallia. Cultures were moistened
with
water
sterilized to pre-
vent
desiccation and, in the
last stages, to favor the antheridia opening and
movement
The
of antherozoids.
culture maintained darkness was opened
in
100 days after sowing. Spores were sown
also in previously
sterilized, natural
organic
soil.
from
Tmax B/W
SEM DSM
94DA.
Results
The
spores bradeorum
of are monolete,
P. bilateral, yellow, and ovate
to
m
PEREZ-GARCIA ET PSEUDOCOLYSIS BRADEORUM
AL.:
221
The
50], spores are chlorophyllous but also contain yellow oily globules. The
thin granulate perine smooth
(Figs. 1, 2) is psilate, to verrucate, or variously
The
ornamented.
laesurae
are short.
Germination was and commenced
of the Gleichenia-tyipe seven days
after
The
sowing. produced
first division a wall parallel to the polar axis of the
spore, cutting off a lateral initial rhizoid cell [Figs. This simple equatorial
3, 4).
form found many members
in of the Polypodiaceae.
is
Several transverse divisions of the initial prothallial cell resulted in a uni-
germ 2-3
seriate filament cells long [each one of these cells vdtli abundant
and The germ and
chloroplasts) a short hyaline rhizoid. filament the rhizoid
A
cell are oriented in opposite directions (Figs. 3,4). large oil globule was
many
apparent in the prothallial cells; this is characteristic in polypodiaceous
ferns.
development bradeorum
Prothallial of Pseudocolysis of the Drynaria-
is
which
type, in the differentiation of the apical meristematic delayed,
cell
is
and generally the prothallus develops trichomes along the margin and on both
and
surfaces. Longitudinal transverse divisions of the filament cells result in
a broadly ovate to spatulate prothallial plate at 20-30 days. The plate has two
or three hyaline basal rhizoids and 36-50 prothallial cells with abundant chlo-
An
roplasts. At this time, there no evident meristematic zone. obconic mer-
is
an one
istematic cell later differentiated as a result of oblique division of of
is
the marginal cells from the distal end of the plate. Young prothalli become
and by
cordiform-spatulate, the meristematic cell replaced a pluricellular
is
meristem At an inconspicuous cushion
differentiated,
(Figs. this stage, is
5, 6).
the prothalli are glabrous, and the old spore wall is retained on the basal cell.
In the D/ynaria-type development the meristematic cell not very active in
is
growth and widening of the plate, as in most Polypodiaceae [Figs. 8
it is 7,
and
9].
advanced development, 70-105 days, gametophytes are cordi-
In stages of
form-spatulate, with a central shallow meristematic zone, symmetrical wings,
abundant reddish brown basal rhizoids, and an inconspicuous cushion
short,
(Figs. 10, 11).
Antheridia appeared 180 days after sowing; they are globose with three cells,
R
a basal fimnel cell, a ring cell, and an opercular cell. In bradeorum, anther-
and
dehiscence accomplished by the detachment of the opercular cell
idial
is
androgenic Archegonia
the opening of a pore in the wall of the cell (Fig. 12).
appear on the medial basal part of the cushion 130-140 days after sowing. The
archegonial necks are slender and oriented toward the base of the prothallus,
neck ma-
with rows 5-6 per row, plus a binucleate cell at
four of cells
cells,
turity
(Fig. 13).
months
The sporophyte appeared about seven after spore
of the
leaf
first
with small whitish hyaline scales the base,
germination; the petiole short at
is
The
the lamina simple and entire, with smooth margins (Fig 14]. venation
is
on
open dichotomous. Anomocytic stomata were observed the abaxial sur-
is
Sporophytes only developed on the natural organic soil cul-
face
(Figs. 15).
tures.
222 AMERICAN FERN JOURNAL: VOLUME NUMBER
91 4 (2001)
jw:^-zrA3.
s.
V
f, •
f.
^msm
r
-^ja
rh-A
® ®
^•*
V^
r*
:
B*i
*!*
>i
.^*
i»T
.
^^V
* --
I
®
®
®
f i
-T
-J _ J -bT
/*»*
i P?^
l\
an
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\\ --SI
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•tit-
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f- ®
Figs. 1-9. Germination and development
prothallial of Pseudocolysis bradeorum
(Rosenst.] D.
L.
SEM
Gomez.
of spore. Spore with yellow
1. 2. oil globule. 3-4. Germination Young
initiation. 5-6.
prothallial plate. 7-8. Spatulate gametophytes.
Spatulate gametophyte with
9. antheridia. Scale
= =
1-6
bars in figs. 15 ^,m, in figs. 7-9 400 \xm.
PEREZ
223
^F -
r .- ^-1 "hh Jl v^ .. :4-^ ,"'
pm
i -
I
vt
)
^^ik
^.<
®
\
4
,%'
*
' LX
f
J. i
1^
t
'Jl
s
n
/
K.
7^
\
®
r^U JK-
#
^5?2
vp
>
Figs. 10-16. Morphology of adult prothallus and sex organs oi Pseudocolysis bradeorum (Rosenst.)
L. D. Gomez. 10. Cordate gametophyte with archegonium. 11. Pluricellular meristem. 12. Antlieridia.
13. Neck of archegonium. 14. Sporophyte. 15. Anomocytic stomata. 16. Gametophyte showing veg-
= =
^m, and
etative propagation. Scale bars in 10, 14 400 in figs. 11, 12, 13, 15 16 15 |xm.
figs.
gemmae
we on margins
After 71 days, observed the formation of the of
When gemmae
young gametophytes detached, these small main-
(Fig. 16).
tained the capacity to grow and developed into adult gametophytes of typical
same
and The development followed he pattern observed
structure sexuality.
in except adult gametophytes were cordate.
agar, that