The Young Gametophyte of Phylloglossum (Lycopodiaceae)

7 Pages·1992·4.1 MB·

by  Dean P Whittier

Annals of the
732
Garden
Missouri Botanical
i
— ——
—
FFiIgGuUrReEsS 1--v9.. Ssppoorreess aanndd ggaammeettoopphhyytteess of rPhhyylllloogglloossssuumm ddrruummmmoonnddiiii.. GUeerrmmiinnaatuinngg ^n"^^^
1 o! SSppoorreess.. 22..
1I.
m
.
are
3. Two-celled gametophyte formed by oblique wall. Most of the and of the proximal cell
— distal cell less
formed
ssDpoorree ccooaatt. 4. Three-celled gametophyte with spore coat attached Arrow indicates the wall
^ to distal cell.
1..
..:„!_„ „f.i. 1_. ithout
second cell division of the gametophyte, which occurred -5. gametophyte
in the proximal Three-celled
cell.
partially
of
umulated urface
Dis^*'
gametophyte 6.
thickened wall of distal cell. Large arrow indicates cell wall formed by second division of »• ^ _
cell
Xof ce
cad cell
of multicellular gametophyte. Arrow waU of ddiissttaall -
indicates mucilage secretion by thickened portion of the
—
g^^
Young
Small
7. multicellular gametophyte. Arrow
indicates mucilage and 9.
thickened wall of distal cell. 8,
young
rhizoids
30 ^m.
Th WaU ceU
differentiation of the distal
^^^|^^^
The
oduced
apparent at the 3 -celled stage (Fig. 5). ^
perpendicular
to the original wall formed by the portion of wall thickens (Figs
this
o
The
either
distal cell does thickening,
in association with the wall
more niu
of
divisions
(Figs. 6, secretion
7). or adjacent there a
it to is
it,
1
&
Number 3 Braggins 733
Whittier
Volume
79,
Young Gametophyte
Phylloglossum
of
1992
«f«r^^^iftn< -I*
•
F
—
Figures
.-
^^~18. Gametophytes oi Phylloglossum drummondii. 10. Globular gametophytes. 1x1*.. S^m-a»l«l» ver-
ir""^ —
auy
tic
oriented
cylindrical gametophyte young Gametophyte with globular base and apical cylindrical
with rhizoids. 1 2.
medium
(near
^^' ^y'^"^'*i^al gametophyte with bend. New apical growth parallel to surface of nutrie—nt
K
ftcal
^^^^ ^^^ oriented New growth away from gravity. 14. Cylindrical
vertically in the horizontal culture tube.
I
^Ea^rmreftophyte
away from source
taken from gametophyte and new growth
reorientation experiment showing bending of
gravity^
15, Young portion without
— photosynthetic gametophyte. Arrow indicates area at base of c:yylliinnddrriiccaall
^nw"
orophyU. Arrow
16, Young New formed Ught hnaavviinngg aa rroouugghh ssuurnfiaicuec..
photosynthetic gametophyte. tissue in
it
^
^ ^"'^ ^-^-.^^».vt-.« w,rh
^^^ ^" gametophytes with
r^''''' Young photosynthetic
^* of cylindrical portion without chlorophyll.- 1177,, 18.
hori^^^'
•^ntal
mm.
orientation
of the noncylindrical. new growth. All scale bars 1
I
aginous
material remains attached to the
5-7; smaU The spore coat often
(Figs. arrows indicate
Thp li^
•muci"l-a&g-e/)•. Tthnee mm>>uu,,cc.^UU:iaageee ^_' -.i i - _» j- ._i ceUii u..n„tti;ll »tVn.eo gtarmaemiputpnnnh;v'tiec .h.aa--? a<xt..t<.a.i..n^ ed a
ssttaaiinnss with alcian blue at distal
f^
2.6 usuaUy gets caught
3%
(0. 7o alcian blue which macroscopic size (Figs. 8, 9). It
in acetic acid),
^monstrates and deforms the surface
that on enlarging cell
contains an mucopoly- the di<=tal
acid
it
Tu^ notch or
•^ccharidi* ppears as a
r .u- A^fn.rwY^\iv nnrM>ar<;
11
734 Annals
the
of
Garden
Missouri Botanical
indentation in the wall where one edge of the spore not produce gametangia in the dark. Cylindrical
coat or was attached 4-7). gametophytes were moved
(Figs.
IS into the
light to deter-
undergo mine whether
photosynthetic gametophytes can
be
tional divisions to give rise to a small multicellular raised in culture and whether they become
will
gametophyte The
young
(Fig, gametophytes sexually mature.
7).
become
more
globular with divisions and ceU growth The gametophytes moved
cylindrical
into the
(Figs. 8-10), At this stage rhizoids begin to develop. light become green (Figs. 15, 16). New growth
the gametophytes
If continue grow from
to in the the apical region develops chlorophyll
first,
become
dark, they
variously sized globular ga- and this region often darker green than older
is
metophytes. However,
they
eventually develop into portions of the gametophyte. Chlorophyll devel-
cylindrical gametophytes
(Figs. 11, 12), which grow opment in the original cylindrical portions of these
up and away
from
the surface of the nutrient gametophytes often slower. Sometimes the basal
is
medium. The
diameter
of the cylindrical gameto- regions of these gametophytes never turn green
phytes appears be
to related to the size of the (Figs. 15, 16; arrows indicate nongreen basal re-
globular gametophyte
from which
they develop
be- gion).
cause
larger globular gametophytes New
give rise to growth of the gametophytes In light is not
thicker cylindrical gametophytes. The A
diameter 15-18) broader
of cylindrical (Figs. or vertical.
the globular gametophyte
can be larger than the apical meristem replaces the conical apex of the
diameter
of the cylindrical growth (Fig. 12). As cylindrical gametophyte. This broad apex forms a
long
as the
cylindrical gametophytes grow more
in the green, thickened gametophyte that grows or
dark, they
continue grow The
to vertically. ga- less horizontally perpendicular to the incident
(i.e.,
metophytes
tend
decrease
to in diameter as the light). Besides the growth in length, the subapical
apices get further from the nutrient medium. With regions of the green gametophyte widen and thick-
some
very long and narrow
gametophytes the api- en (Fig. 16). The thickest part the median dorsal
is
growth
cal appears
to be very slow or to stop. This region, making these areas hemispherical in cross
appears
related to the distances that carbohydrate The new growth rough
section. surface of the is
has to move from medium
the nutrient through compared to the smooth surface of the white cy-
the gametophytes
to the apical growing gameto-
points. gametophytes. Although these
lindrical
The
growth
of all the cylindrical gametophytes phytes are about one and a half years old, they
vertically and away from
the surface of the nutrient are immature and do not bear gametangia.
still
medium
suggested
a negatively Even axe
gravitropic re- though the gametophytes grow slowly in
To
sponse.
test this, cultures with gametophytes toward
nic culture" at this time they exhibit a shift
growing
were on
vertically
laid their sides (hori- their mature morphology.
and
zontally) maintained dark
in the for five months.
The
apices of these gametophytes
responded
to the
Discussion
positional change by bending
at right angles and
growing away
from
the source of gravity The Phylloglossum game-
(Figs. 13,
best description of
The
gametophyte
14). in Figure 13 was photo- tophytes by Thomas He described the
(1901).
—
is
—
tube
tubercle
gametophytes primary
as having a
experiment. The
direction of the new growth shaped
was
upright and an irregularly
cylindrical shaft,
^'^'^
parallel to the surface of the nutrient medium
which
photosynthetic crown bearing gai
had
a vertical orientation
in the horizontal ^amf
culture voun^est
described
^le
tube The
gametophyte crown
13).
in Figure
14,
which shows same
the
reorientation of th
to gravity, was had He length
not developed. noted that the
m
tube
and tnai
Shalt varied on mature gametophytes
Gametophytes
in the reorientation
test changed crown
be tuberce
primary
the ends near and includmg the
gametophytes
with ha
right angle
bends. These gametophytes
ga- were without These
chlorophyll.
metophytes
demonstrate
that the developed
cylindrical ga- a mycorrhizal fungus was best
that
metophytes
growing
in the dark suggested
are Thomas
negatively
their lower
portions. Finally,
Me
gravitropic.
begm
"may
that Phylloglossum gametophytes
Mature
gametophytes
of Phylloglossum
are endophytic
de- as a saprophyte dependent on an
scribed as being green and
photosynthetic (Thomas
gus."
m
axenic
Thi
ture maintain
their
white, cylindrical hab.t and do germination
axenic culture account of
the
is first
r _-
&
Number 3 Whittier Braggins 735
Volume
79,
Young Gametophyte
Phylloglossum
of
1992
L
Table Qiaracter comparisons of Phylloglossum and subgenus Lepidotis of Lycopodium,
Character Phylloglossum Lepidotis
Basal mucilage duct in sporophyll absent present
Epidermal walls of sporangium sinuate straight unlignified
lignified
Spore ornamentation
foveolate rugulate
Spore germination
dark
light
Nutrition of young gametophyte mycorrhizal photosynthetic
Position of mature gametophyte
surficial surficial
Nutrition of mature gametophyte photosynthetic photosynthetic
Photosynthetic lobes on gametophyte absent present
Multicellular uniseriate hairs on gametophyte absent absent
Embryo
type
large foot small foot
and
early gametophyte
growth
for Phylloglossum. Thus, morphological changes the gametophytes
to
rrior
to
this report, the youngest gametophytes inthelight occur both the and axenic culture,
in soil
described were
those with a primary tubercle and The development of these gametophytes in cul-
cylindrical
shaft (Thomas,
1901). Bertrand(1885), gametophytes from na-
ture similar to that for
is
Sampson
(1916b), and HoUoway
(1935) faUed Growth gametophytes up through the
to ture. of the
germinate
the spores of Phylloglossum. An by
earlier soil to the light has to be simulated in culture
and
very
brief report by
Crie (1883) reported the moving the gametophytes from the dark to the
germination
of
these
spores but did not describe Also gametophytes in culture are dependent
light.
the
early
stages
of gametophyte He medium
development. on sugar the nutrient for their devel-
in
report
mature,
'^^ white, bulbous gametophytes opment and on mycorrhizal fungus nec-
not the
be
essary gametophyte growth in nature.
for
Because
^^^^^^^' the gametophytes he on growth and development of
de- Observations the
^k^j
" do
not
the can
fit descriptions of gametophytes Phylloglossum gametophytes axenic culture
in
(Thomas,
1901; Sampson, 1916b; how they grow nature. Phyllo-
help explain in
to
Holko,wwa"^y*,"'^
1935)
or from and
axenic culture, we choose glossum spores germinate only in the dark,
to
disregard
the
Crie gametophytes
report. the white cylindrical stage of these
e
^-celled gametophyte of Phylloglossum negatively gravitropic. would appear that the
It
is
nsists
of a
proximal and The being covered
distal cell. distal spores in nature germinate only after
underg
The
by or percolating into the soQ. early de-
soil
pgion
of
ceU waU
its thickens and mud- gametophyte requires a
secretes velopment of the white
3ge- At
maturity
the Be-
distal ceU of these gameto- mycorrhizal fungus for organic nutrition.
its
^^
.^^"^^ similar to the proximal young portion negatively
of cause the white cylindrical is
bL f cell
^^™*^^°P^y^^s, which has a mucilage- gravitropic, grows up through the soil until it
it
coat^'d^
l!'-'" &
^^"
(Melan The Once the apex of the
Whittier, 1989). reaches the surface.
soil
prox
1
^^^ °^ *^^ young Phylloglossum game- gametophyte exposed to the light, its
cylindrical is
tODh""^
P e
with
y and photosyn-
Its thin wall remains meristematic. developmental pattern changes the
rough
divisions crown
of the proximal
ceU, the multi-
^^'
gametophyte
is formed, stage there any possibility for sexual repro-
this
is
^^^^^^'^ gametophyte
forms
early in the duction.
develo
^^ .,— The gametophytes and confusion
^^y^^'^S^ossum gametophytes photosynthetic
in
s«,w^tv.t.,v.^o
^thgeV^L^^"^ o^...^...,. ...
,
ark.
In axenic (Bower, 1886),
PAj/Zo^/ossu/n
culture usually does not about the tuber of
it
ome
large having
before many Phylloglossum as
the initiation of the cylindrical led to consider
metophyte. mem-
However, Lycopodium cernuum or other
some
in cases the globular with
affinities
Thomas
(1901)
^^^ are larger than the diameter of the bers of subgenus Lepidotis^. Also,
cyl^d
growths
rLJ^T^' that arise from them. In these
cases
0-— Although recognition
me cernuum.
^'wpiijf ics ni aescripuon oi a similar to that of L.
p_-
ry
tubercle
and
by tuber
cylindrical shaft described
'nomas \
r
1916a;
(igoT\ cernuum (Sampson,
•«
i moved. - protocorm of L.
"^/- '-'Smetophytes to ulumi- to the
laterf
number propose J
J of
reduced the
appear game- Osbom, 191 has
*"'"®^ to be the 9)
initiating
toph
^,^ and subgenus
Phylloglossum
^''o^n between
as Thomas
described by (1901). similarities
736 Annals
the
of
Garden
Missouri Botanical
Phylloglossum W. A
Lepidotis, generally considered Harris, 1955. manual
is F. of the spores New
of
New
Zealand
Pteridophyta. Zealand
Dept.
affinities Sci. In
Government
dustrial Res. Bull. 116, Printer, Wel-
way, 1935; Hackney, 1950; Bruce, 1976a).
lington.
In an examine more any
eff"ort to closely possible
Hollow
AY, E. 1935. The gametophyte
of Phyllo-
J.
similarities between Phylloglossum and subgenus
glossum drummondii. Ann. (London) 513-
Bot. 49;
Lepidotis, a of sporophytic and gametophytic 518.
list
Knox, M. The
characters are presented Table The E. 1950. spores of Lycopodium, Phyl-
in sporo-
1.
loglossum, Selaginella and and
Isoetes their value
phytic characters, which are few, do not include
in the study of microfossils of Palaeozoic age. Trans.
those used Phylloglossum
to classify the Ly-
in
209-357.
Bot. Soc. Edinburgh 35:
copodiaceae or those altered by the reduced habit Melan, M. a. & D. P, Whittier. 1989. Character-
of Phylloglossum, Although a more exhaustive ization of mucilage on the proximal cells of young
gametophytes of Botrychium dissectum (Ophioglos-
comparison
awaits a complete
description of the
Amer. 1006-1014.
saceae). Bot. 76:
mature gametophyte J.
of Phylloglossum, appears
it A
0LLGrARD''B 1987. revised classification of the Ly-
that Phylloglossum and subgenus Lepidotis have
153-178.
copodiaceae Opera Bot. 92:
lat.
s.
few similarities (Table 1). The number of differ- OSBORN. T. G. B. 1919. Some observations on the tuber
485-516.
ences between Phylloglossum and oi Phylloglossum. Ann. Bot. (London) 33:
subgenus Lep-
W.
ROTHMALER, Feddes
1944. Pteridophyten-studicn.
I.
idotis suggests that a reexamination
of past pro-
55-82.
Repert. Spec. Nov. Regni Veg. 54:
posals of between them
affinities
in order.
is
Sampson, K. 1916a. The morphology Pftj//og/o55um
of
315-
drummondii Kunze. Ann. (London) 30:
Bot.
331.
P/tj/^/oisu/n
1916b. Note on a sporeling of
Literature
Cited
.
attached to a prothallus. Ann. Bot. (London) 30:
Bertrand,
C. E. 1885. Phylloglossum Drummondii. 605-607.
Arch. Bot. N. France 70-223. & 1959.
2: Sheat, D. E. G., B. H. Fletcher H. G. Street.
BoiviN, B. 1950. The problem of generic segregates VIIL The
in Studies on the growth of excised roots.
the form-genus Lycopodium. Amer. 32-
Fern 40: growth of excised tomato roots supplied with various
J.
41. New ilo-
inorganic sources of nitrogen. Phytol. 58:
Bower, On
F. O. 1886. the development and mor-
154.
phology of Phylloglossum drummondii. Thomas, W. account of the
Philos. A. 1901. Preliminary
P.
Trans. 176, 665-678. Lon-
II: Roy. Soc.
prothallium o{ Phylloglossum. Proc.
&
Breckon,
G. R. H. Falk. 1974.
External spore don 285-291.
J. 69:
morphology and taxonomic of Phylloglos- Wagner, W. & M. 1992. Generic
affinities H., Jr. Beitel.
J.
sum
drummondii Kunze
(Lycopodiaceae). Amer. modern North American Lycopodi-
J. classification of
481-485.
Bot. 61: 676-686.
aceae. Ann. Missouri Bot. Gard. 79:
Bruce,
G. 1976a. Development and 5o^rjc/uw'"
J. distribution of Whittier, D. 1972. Gametophytes of
P.
mucilage canals Lycopodium. Amer. ^az.
in Bot. 63: Bot.
J. dissectum as grown in sterile culture.
481-491.
336-339.
— (Crawfordsville) 133:
1976b. Gametophytes and subgeneric con- Lycopodium digi-^
. 1981. Gametophytes of
.
cepts Lycopodium. Amer. 919-924,
in Bot. 63:
J. latum (formerly complanatum var flabellifc
L.
.
Bruchmann,
H. 1898. Uber
und
die Prothallien (Crawfordsville)
die as grown in axenic culture. Bot. Gaz.
Keimpflanzen mehrerer
europaischer
Lycopodien.
519-524,
142:
Gotha. & of
The induction
1960.
T. A. Steeves.
CRife, L. 1883. Essai sur
la flore primordiale. Octave Bot.
apogamy Canad.
in the bracken fern. J-
Doin,
Paris.
925-930.
Hackney, M. A
F. V. 1950. review of and contribution Wilce, L General spo^^
H. 1972. Lycopod spores,
J.
knowledge
to of Phylloglossum drummondii Lycopodium.
Kunze.
patterns and segregates of
the generic
New
Proc. Linn. Soc. South Walps 7S- ^'^-i ^9
1 Amer. Fern 65-79.
62:
J.
Number
207-445
2, pp. of the Annals of the Missouri Botanical
was May
published on
1992.
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CONTENTS
The A &
Symposium
Lycopsida:
William DiMichele Skog
A. Judith
E.
Phylogenetic
Relationships of the Zosterophylls and Lycopsids: Evidence from Mor-
phology, Paleoecology, and Methods
Cladistic of Inference Patricia
G.
Gensel
..„ 450
Thoughts on
the Early Lycopsids and Zosterophylls Francis M. Hueher 474
Experimental
Cladistic Analysis of Anatomically Preserved Arborescent Lycopsids
from
the Carboniferous An
of Euramerica: Essay on
Paleobotanical
&
Phylogenetics Richard M. Bateman,
William A. DiMichele Debra
A.
Willard
500
Arbor
L
Swamps &
boniferous of Euramerica Tom
Phillips William A.
DiMichele
560
Evolution
of Isoetalean Lycopsids Kathleen Pigg 589
B.
&
Habitat, Evolution, and
Speciation in Isoetes W. Carl Taylor R. James
Hickey
613
Paleozoic Herbaceous
Lycopsids and the Beginnings of Extant Lycopodium sens.
lat.
and
Selaginella sens. Barry Thomas 623
A.
lat.
Developmental
Problems
Selaginella
in (Selaginellaceae) an Evolutionary
in
Context
Terry R. Webster 632
The &
Mesozoic Herbaceous / 648
Lycopsids Skog
E. R. Hill
C.
Modern
Warren
&
ner, Joseph M. 676
Jr. Beitel
Neotropical Lycopodiaceae— An 687
Overview Benjamin 0llgaard
Cytological Problems 718
in Ljco/)oofiu/o Wagner
sens. Florence
lat. S.
The Young &
Gametophyte
of Phylloglossum Dean
(Lycopodiaceae) Whittier
P.
John
Braggins 730
E.
Cover
illuslration.
Sparcxi.
viUosa (Burman
Coldblatt. by M.
f.) Branch.
L.