Spore bearing structure.
May be eusporangiate or leptosporangiate.
Homo- or Heterosporous.
Terminal/ Lateral/ aggregated into specialized structures.
Sporangia are borne on unmodified foliage or on specialized fertile leaves.
Thick or thin walled.
Either terminal on the leaf segments but more commonly they are on the frond, marginally or superficially and rarely in specialized structures, the sporocarps.
Sporangia either single or form sori or fuse to form synangia.
The sporangia in higher ferns are aggregated in groups known as sori.
Sori may be circular to linear.
Positioned along the margin or towards the midvein (costa), surficial or in a groove or channel, etc.
In some cases, the developing sori are protected by a recurved leaf margin (false indusium), a covering of deciduous scales, or a more permanent small flap of tissue, the indusium.
Indusia vary greatly in shape, size, texture and persistence, ranging from umbrella-shaped to globose to linear.
Sori may be variable in size, the larger ones result due to fusion, the coenosori.
The protion of the leaf surface bearing sporangia is termed as receptacle, it may be an elevated surface or a flat elongate structure.
Sporangia originate from the superficial cells of the receptacle.
Tassel is regarded as primitive and acrostichoid condition as advanced.
Hubble Asteroid Hunter III. Physical properties of newly found asteroids
Soral & Sporangial Characters in pteridophytes.pptx
1. Soral and Sporangial
Characters in Pteridophytes
- A comparative account
(https://th.bing.com/th/id/R.f407870c5eb7f427fe075c90de1e510d?rik=mHaijxcSeHv9fQ&riu=http%3a%2f%2fi1.treknature.com%2fphotos%2f482%2fff1.jpg&ehk=swnMZ%2fQnONqiBzoH0ckmgVsJzbudAKzbHSnKmZN%2fhn0%3d&risl=&pid=ImgRaw&r=0)
3. Sporangia
•Spore bearing structure.
•May be eusporangiate or leptosporangiate.
•Homo- or Heterosporous.
•Terminal/ Lateral/ aggregated into
specialized structures.
•Specialized structures may be sporophyll/
strobili/sporocarp/sorus.
(https://i.pinimg.com/originals/98/92/bc/9892bc880bc70ff9c5cc81b22f9312f8.jpg)
4. Sporangia
• Large sporangia are found in
Ophioglossales & Marattiales.
• The more primitive families of ferns:
Osmundaceae, Gleicheniaceae &
Schizaeaceae also have large sporangia but
the walls are thin and annulus unspecialized.
• The annulus is presumed to have originated
by gradual differentiation within the surface
layer until complete dimorphism of all types
resulted.
• Sporangia of Actinostachys, Schizaea,
Stromatopteris and Osmunda show
intergradation between surface cells and
annular cells.
(http://cupac.bh.cornell.edu/cupac_images/2013_02_08/0000031.21.tif.jpg)
(http://cupac.bh.cornell.edu/cupac_images/2013_02_09/0000027.9.tif.j
pg)
6. Dehiscence of a Sporangium
• In the eusporangiate ferns, dehiscence is by a slit and is fundamentally
longitudinal in all cases.
• In the transitional family, Osmundaceae, Osmunda & Todea, The
annulus is located near the tip but on to one side and forms a cleft
that runs over the top of sporangium and down the opposite side. The
sporangia opens like a clam.
• In the leptosporangiate ferns,
qthe annulus may form a cap at the distal end of sporangium
as in Lygodium longitudinal dehiscence
(https://th.bing.com/th/id/R.847257aa97cf9d9599adf6f434d58c6a?rik=KTZtRQSDMZAL9A&riu=http%3a%2f%2fpalaeos.com%2fplants%2fglossary%2fimages%2fAnnulusLygodium.jpg&ehk=LLST2hvjWErgWH0stEdFKo3%2bQAnpvMYcL%2b%2fSTVcIpTE%3d&risl=&pid=Im
gRaw&r=0)
7. q be obliquely placed as
in Gleichenia, Hymenophyllum & Loxsoma oblique dehiscence
qRun over the top of sporangium in line with stalk i.e., in vertical position
as in Leptochilus vertical dehiscence.
8. Development of
sporangia
Eusporangiate Leptosporangiate Transitional type
https://th.bing.com/th/id/R.74b63b79d3cc1977102904640511417c?rik=pgzmWB5BMaEIrA&riu=http%3a%2f%2fbotit.botany.wisc.edu%2fResources%2fSystematics%2fLycopodiophyta%2fLycopodiales%2fLycopodiaceae%2fLycopodium%2fUndetermined+species%2fSporangium+ls+M
C.jpg&ehk=bCKBSv3EKmW%2bhiPYbci%2f%2fuFsElXOlEsqhnGCq4EGNcI%3d&risl=&pid=ImgRaw&r=0, https://media.gettyimages.com/photos/pteris-50x-at-35mm-also-shows-spores-and-annulus-picture-
id139802925, https://live.staticflickr.com/2692/4382051069_eae4a41938_b.jpg)
22. Sporangia in Rhynia
• Borne singly on the apices of some aerial branches.
• Each sporangium being oval or slightly cylindrical structure with a
little greater diameter than that of aerial branch on which it is
developed.
• They were 12 mm long and 4 mm in breadth in R. major and 4
mm long and 1 mm broad in R. gwynne-vaughani.
• Five cells thick wall.
• 1 cell thick cuticularized epidermis was followed by 3 cells thick
middle layers of thin walled cells.
• The inner-most layer was 1 cell thick tapetum.
• The wall was surrounding a spacious sporangial cavity.
• Columella absent.
• Homosporous with numerous spores.
• No special mechanism of sporangium dehiscence.
• The liberation of spores by disintegration of the sporangial wall.
23. Synangia in Psilotum
• Eusporangiate development.
• Synangium- fusion of 3 sporangia.
• Borne on the axils of small leaf-like
appendages.
• Neither its outermost sporogenous cell
nor the innermost jacket cells develop
into the tapetum.
• The spores formed in the tetrads are
colourless and reniform.
• Homosporous.
(https://live.staticflickr.com/3094/3117458944_65d8090a24_n.jpg)
(https://botit.botany.wisc.edu/Resources/Botany/Pteridophytes/Psilophytes/Psilotum/Synangium%20MC.jpg)
24. Sporangium in
Protolepidodendron
• On the middle part of the stem
sporophylls are laxly borne intermixed
with vegetative leaves.
• The sporophylls resemble the vegetative
leaves with bifurcate tips and bear single
sporangia on the adaxial face is little
above the bases.
• No vascular connection leading to the
sporangia.
25. Sporangial Characters in
Lycopodium
• Found singly on fertile leaves known as sporophylls.
• Sporophylls bearing sporangia densely aggregated along stem
to form either branched or unbranched called strobilus.
• Strobili are terminal.
• Reniform, orange to yellow.
• Unseptate sac-like structure with a narrow or massive stalk.
• Homosporous.
• Eusporangiate development.
• Dehiscence by transverse rupture of sporangium apex along
the line of stomium. (https://asset.library.wisc.edu/1711.dl/STGCBU6ULSV2I8X/M/h1380-e86e1.jpg)
(https://www.researchgate.net/profile/Alberto_Diego-
Taboada/publication/260875436/figure/fig1/AS:296782396444675@1447769845877/Scannin
g-electron-microscopy-SEM-picture-of-Lycopodium-clavatum-spores-showing.png)
26. Sporangial
Characters in Isoetes
• Heterosporous.
• Largest among all known extant spore-
producing plants.
• Highest spore output.
• 50-300 megaspores/ megasporangium.
• 150,000-1000,000
microspores/microsporangium.
• Sporangium traversed irregularly and
incompletely by plate-like trabeculae.
• Trabeculae surrounded by tapetum.
• Microspores elongate, 45 microns long.
• Megaspores 250-900 microns in diameter. (https://th.bing.com/th/id/OIP.pb3eLzKrqAdGbLJVstdVcQAAA
A?pid=ImgDet&rs=1)
(http://idtools.org/id/
aquariumplants/Aqua
rium_&_Pond_Plants_
of_the_World/key/Aq
uarium_&_Pond_Plan
ts/Media/Images/Isoe
tes-melanopoda-H-
wilson_lge.jpg)
(https://www.researchgate.net/profile/Paulo-
Labiak/publication/272214264/figure/fig5/AS:670531535765526@1536878584817/SEM-images-of-the-spores-
of-Isoetes-maxima-Pereira-et-al-631-UPCB-I-pedersenii.ppm)
27. • All leaves are potential
sporophylls.
• Sporophylls are ligulate.
• Eusporangiate.
• In microsporangia the sporocytes
differentiate after the formation of
trabeculae, while in
megasporangia the sporocytes
differentiate before the formation
of trabeculae.
• No dehiscence mechanism.
• Spores are released only after the
death and decay of the sporangia.
(https://newfs.s3.amazonaws.com/taxon-images-1000s1000/Isoetaceae/isoetes-echinospora-sc-blegler.jpg)
28. Sporangial Characters in Selaginella
• Sporangia are produced on the axils of ligulated
leaves called sporophylls.
• The sporophylls are compactly arranged to form
cones or strobili.
• Heterosporous.
• Eusporangiate.
• Sporangia reniform to ovoid with a short stalk.
• In microsporangia, all sporocytes except 10-
20% from form microspores.
• Each sporangium has 2 layered wall, a layer of
tapetum and sporogenous tissue.
• Spores are with a prominent triradiate mark and
characteristic ornamentation.
(http://1.bp.blogspot.com/_mFm838BFqBE/TQFEUcH2-7I/AAAAAAAAAUY/Qq0xw6_OtX8/s1600/esporangios-selaginella.jpg)
(https://tse1.mm.bing.net/th?id=OIP.eodQ88o23qLQAGIEsDz
W9wHaFf&pid=Api&P=0&w=255&h=189)
(https://www.researchgate.net/profile/Ivan_Valdespino2/pu
blication/290156962/figure/download/fig3/AS:56392429075
2513@1511461436311/Selaginella-nanuzae-Valdespino-A-
Megaspore-proximal-face-B-Close-up-of-megaspore.png)
(http://www.sciencephoto.com/image/16797/large/B400009
8-Selaginella_moss_strobilus,_LM-SPL.jpg)
29. S. selaginoides Basal sporangium are non-
functional.
S. sulcata One meagaspore is produced
S. rupestris Two megaspores are
produced.
S. helvetica Megasporangia at the base &
microsporangia at the distal
region.
S. helvetica S. rupestris
S. selaginoides: discharge of megaspores by
"compression & slingshot ejection"
(https://bio.libretexts.org/@api/deki/files/29958/InkedSelagi
nella_strobilus_LS_high_LI.jpg?revision=1&size=bestfit&widt
h=456&height=685)
30. S. oregana Megasporangia on one side
and microsporangia on the
other side.
S. kraussiana One lower megasporangium
and upper microsporangium.
S. inequifolia Microsporangia on the left
side and microsporangia on
the right side.
S. cuspidata The apex may continue
vegetative growth beyond
the formed strobilus and
thus produces a shoot
with sporophylls and foliage
leaves in alternating
zones along the stem.
S. oregana
S. inequifolia
S. kraussiana
31. Sporangium of Hyenia
• The sporangiophores were forked and
the tips were reflexed which in turn
terminated in two or three pendent
sporangia.
33. Sporangium in
Calamites
• Several types of strobili have been
found on the fossil Calamitean type of
plant body.
• These have been assigned to the organ
genera: Palaeostachya, Calaamocarpon,
and Calamostachya.
• In all strobili, whorls of peltate
sporophylls alternated with whorls of
lanceolate bracts.
• Strobili were either homo-
/heterosporous.
34. Sporangium in
Equisetum
• Reflexed sporangia are produced on sporangiophores which
are aggregated into definite cones or strobili at the apex of
fertile shoot.
• Homosporous.
• Eusporangiate.
• Strobilus is composed of
sporangiophores: an axis with whorls of stalked, peltate
structures.
Immediately below the whorls of sporangiophores, the axis
bears a small or large ring like outgrowth, the annulus.
• Each sporangiophore is umbrella like in shape.
• 5-10 sporangia on the underside of each sporangiophore disc.
Annulus
Vestigeal remainder of
Calamitean bracts.
Sporangiophoric in nature, since
it sometimes bear small
sporangia.
(https://live.staticflickr.com/895/2805
2498497_d2ba692831_b.jpg)
(https://th.bing.com/th
/id/R.3c58d8cb1fbca45
d006b2fedb525e7df?ri
k=cPxiTXHzy41gtA&riu=
http%3a%2f%2fwww.pl
antsystematics.org%2fu
sers%2fkcn2%2f4_22_0
4%2fUpEquisetum%2fE
qui_arvense6.jpg&ehk=
WUYQon%2bouoVQtxQ
OYD8v%2fIlmfj9Xh1sdb
8AH7ihL4Ms%3d&risl=
&pid=ImgRaw&r=0)
35. • Sporangia: slender stalk
bearing a peltate and
hexagonal disc on the under
surface of which there
are elongate,
cylindrical hanging sporangia.
• Each sporangium is an
elongated, sac-like sructure
attached to the inner side of
stalked, sheild-shaped
sporangiophore.
• At first, the wall is 3-4 layered
but when mature 2 jacket
layers remain.
(https://c2.staticflickr.com/6/5318/5883581480_32215
18fce_z.jpg)
(https://upload.wikimedia.org/wikipedia/commons/thumb/7/77/Equisetum_arvense_spora
ngia.jpg/360px-Equisetum_arvense_sporangia.jpg)
37. Spores of Equisetum
• Spores are globular/ spherical in shape.
• Thick wall with 4 concentric layers:
Innermost cellulosic, delicate intine.
Exine
Middle layer
Thick epispore
• Intine and exine are products of spore-protoplast.
• Middle layer & Epispore are formed around the intine
& exine by the activity of the periplasmodium.
• Epispore splits to 4 spirally arranged ribbon like bands
called elaters, flattened at the tips.
• The bands are free from the spore wall except for one
common point of attachment.
(https://i.pinimg.com/originals/59/bf/32/59bf326a69b49c18
545b2aca8f3f5ba8.jpg)
(https://www.nirgal.net/microscopie/spore_prele_zstacking_big.jpg)
38. • The elaters are hygroscopic.
• When moist, they are spirally coiled round the spore
and uncoil and become extended as the spore dries
out.
• Elaters are different from those found in bryophytes.
• When the spore matures, elaters uncoil and this
expansion of the elaters help in the dehiscence of
sporangium.
• In shedding, these elaters uncoil, become entangled
with the elaters of other spores and thus the spores
are shed in groups rather than singly.
• The elaters aid in spore dispersal.
• The spores contain chloroplast.
(http://www.missouriplants.com/images/Equisetum_hyemale_spores.jpg)
(https://tse3.mm.bing.net/th?id=OIP.gh3gpsLFP77dMGbyjzzjGAHaFj&pid=Api&P=0&w=215&h=161)
39. • Sporangia are borne on unmodified foliage or on specialized fertile
leaves.
• Thick or thin walled.
• Either terminal on the leaf segments but more commonly they are on
the frond, marginally or superficially and rarely in specialized
structures, the sporocarps.
• Sporangia either single or form sori or fuse to form synangia.
General Characteristics of Sporangia in
Pteropsida
40. Sorus
• The sporangia in higher ferns are aggregated
in groups known as sori.
• Sori may be circular to linear.
• Positioned along the margin or towards the
midvein (costa), surficial or in a groove or
channel, etc.
• In some cases, the developing sori are
protected by a recurved leaf margin (false
indusium), a covering of deciduous scales, or
a more permanent small flap of tissue, the
indusium.
(https://rbg-web2.rbge.org.uk/thaiferns/factsheets/data/images/Adiantum_stenochlamys/thumbs/Adiantum_stenochlamys_Middleton_et_al_5221_PK_8269_sml.JPG)
41. Sorus
• Indusia vary greatly in shape,
size, texture and persistence,
ranging from umbrella-shaped
to globose to linear.
• Sori may be variable in size,
the larger ones result due to
fusion, the coenosori.
• The protion of the leaf surface
bearing sporangia is termed as
receptacle, it may be an elevated
surface or a flat elongate
structure.
• Sporangia originate from the
superficial cells of the receptacle.
(https://i.pinimg.com/originals/12/cf/a3/12cfa3df749b572051e98ac93e4e99e0.jpg)
42. Coenosori may be broken up
into segments as in Blechnum &
Woodwardia.
(https://live.staticflickr.com/4075/4822598915_b71f54b446_b.jpg)
(https://live.staticflickr.com/130/377901147_39651a875d_b.jpg)
43. Felt & Marginal
Tassels
• If sporangia do not form sori,
they are either scattered over
the lower leaf surface forming a
felt as in Acrostichum or
marginal tassels over the
surface of slender leaf divisions
as in Osmunda or are solitary,
along or close to the margins of
narrow leaf segments as in
Schizea and Anemia.
(https://fernsofmacaronesia.org.uk/wp-
content/uploads/2020/06/Osmunda-regalis-DSC04760B-768x512.jpg)
(https://www.fernsoftheworld.com/wp-
content/uploads/2014/05/Schizaea-
elegans_PPDaniels_1047-1000x640.jpeg)
(https://tse2.mm.bing.net/th?id=OIP.7qilqlhdjEUevrSL21RUpwAAAA&pi
d=Api&P=0&w=206&h=158)
45. Marginal sorus:
• Receptacle and sporangia originate from
the margin of pinnae or pinnule.
• The indusium, if present is a
submarginal outgrowth around receptacle.
• Indusium is funnel shaped or bilipped.
• The two parts of the indusium may be
different, often adaxial flap is thicker and
histologically ressembles the leaf.
• e.g., Hymenophyllum, Dicksonia, Culcita,
Cibotium
(https://c1.staticflickr.com/7/6068/6030902369_0ef8285b5a_b.jpg) (https://live.staticflickr.com/2644/3938743546_5c0e706374_b.jpg)
46. Intramarginal sorus:
• The marginal cells often form a thin
extension or loose meristematic
activity.
• The submarginal cells on abaxial side
of the lamina become meristematic
and form the receptacle or
sporogenous meristem.
• In some cases, a thin abaxial indusial
flap is also formed from the surface
cells near the meristem.
• E.g., Cryptogramma, Pellaea,
Pteridium, Adiantum
(https://tse3.mm.bing.net/th?id=OIP.SapKwbGymNEAdl4ijg2drQAAAA&pid=Api&P=0&w=158&h=158, https://tse1.m
m.bing.net/th?id=OIP.L3yBoGzBjN8opUylrgntawExDM&pid=Api&P=0&w=241&h=161)
47. Superficial/ Abaxial sorus:
• Receptacle is submarginal in origin on
lower surface but the margin of lamina
remains active and continues to add new
tissue shifting the sorus farther away
from margin.
• The indusium, if present, is formed from
superficial cells of receptacle and
overarches it.
• In some forms indusium originates
from the top of receptacle forming a
stalk and a peltate disc.
• e.g., Asplenium, Gleichenia,
Athyrium, Aspidium
(https://live.staticflickr.com/3450/3186416993_1548b1864e_b.jpg, https://wcbotanicalclub.files.wordpress.com/2020/10/asplenium-platyneuron-ebony-spleenwort-sori.jpg?w=1024)
50. Phyletic side of a Sorus
Marginal position.
Migration.
Superficial position
(https://media.istockphoto.com/photos/sword-fern-spores-on-fronds-picture-id1188490560?k=6&m=1188490560&s=612x612&w=0&h=UJzWwjIwhs3RPldd7PdkgvKQPqPH7vR62GE04YmDqHs=)
51. From a tassel to a sorus
• Tassel is regarded as primitive
and acrostichoid condition as
advanced.
• Sori are described to be
derived from the tassel in
three ways:
1. A superficial sorus can be
obtained from the tassel by
the development of a blade
on the axis. This is seen in
Osmunda-Todea series.
(https://s-media-cache-ak0.pinimg.com/736x/cb/14/d8/cb14d8a20f846677117afa111696e702.jpg)
52. 2. A blade with monosporangial sori as in Lygodium can be obtained by a similar expansion as in Osmunda-
Todea series but along the axis of sporangial attachment. The solitary sporangia later form sori by increase in
number. This can be visualized through Lygodium, Mohria and Gleichenia.
3. The tassels can change into marginal sori by the webbing of sterile stalks of fertile tips. This may be
seen in Anemia, Schizaea and Trichomanes series.
(https://www.polyploid.net/swplants/images/Lygodium_jap_3Jul07_2008.jpg, https://tse2.mm.bing.net/th?id=OIP.sFvYFbezciRURkzcuYtOtAHaFN&pid=Api&P=0&w=238&h=167, https://www.kaimaibush.co.nz/ferns/Resources/sticheruscunning1.jpeg)
https://ww.backyardnature.net/n/10/101226ao.jpg, https://tse3.mm.bing.net/th?id=OIP.4u7OKrkpgPGPu-FhfT8cswHaFk&pid=Api&P=0&w=224&h=168, https://live.staticflickr.com/15/18882865_aeeff55055_z.jpg)
53. Protection of a Sorus
• Protection of young sporangia during
ontogeny is provided by circinate
vernation.
• The sporangia on expanded
sporophyll are with or without the
indusium(protective covering).
• The sori
lacking indusia are exindusiate or
naked (Gleicheniaceae).
• In forms having marginal sori, the
indusium might become a cup or
pouch-like structure formed by
extension of the abaxial and
adaxial margins of leaf (Trichomanes,
Davallia, Dicksonia).
Dicksonia
Davallia
Davallia tasmanii
Gleichenia
(http://www.nzflora.info/factsheet/images/Image1YX8large.jpg
)
(https://tse2.mm.bing.net/th?id=OIP.cbF03HezvPVpKPEbFfyO-
gDLEy&pid=Api&P=0&w=99&h=148)
(https://farm3.staticflickr.com/2483/3899270245_8ca08b372f_z.
jpg?zz=1)
(http://farm3.staticflickr.com/2644/3938743546_5c0e706374_z.jp
g?zz=1)
54. • In forms having superficial sori, the
indusium is essentially an outgrowth
of leaf epidermis.
• The indusium is vairable in its form
and extent.
It may be delicate strip attached
along one side only-
Unilateral indusium ( Asplenium).
Reniform and attached at
sinus (Dryopteris).
A peltate disc (Polystichum,
Cyrtomium)
Circular
Horse shoe shaped
Cup-shaped
Dryopteris
Asplenium
Polystichum
(https://delawarewildflow
ers.org/images/110911_13
1150.jpg)
(http://2.bp.blogspot.com/-
x6CBXBX6jtQ/VAsLoiRI0jI/AAAAAAAAbzs/65AMLtVg
ycU/s1600/Dryopteris%2Bfilix%2B-
mas%2B'Euxinensis'%2B%2Bspores%2B06092014%2
B(4).JPG)
(https://tse3.mm
.bing.net/th?id=
OIP.Co53SMSgjZ
H28A-
G9GedaQAAAA&
pid=Api&P=0&w
=255&h=188)
55. • Instances where reflexed leaf
margin provide protection to
sporangia are refered as false
indusia.
• The indusium in superficial sori
can be seen to have
undergone a further change.
The point of
attachment migrated from
margin to the centre of
receptacle and it became a
radially symmetrical structure.
• From a radially symmetrical
indusium, an elongate
lateral indusium is derived by
the extension of the area of
attachment along on side.
This is seen in Aspidium-
Athyrium-Asplenium series.
56. Maturation of Sporangia within a Sorus
Simple Sorus:
• A sorus in which all of the sporangia originate, grow and mature at the same time.
• Forms showing such conditions are grouped together as 'Simplices' but are not
necessarily closely related.
• Primitive
• Found in ancient fossil ferns
Gradate Sorus:
• If sporangia develop over a period of time with oldest sporangia in the centre and
successively younger sporangia towards the base.
• Forms showing it are grouped together as 'Gradate'.
Mixed Sorus:
• When sorus shows an intermingling of sporangia of different ages it is a mixed sorus.
• Condition is termed 'Mixtae'.
• More highly specialized and evolved families have this mode of soral development.
57. Sporangia in
Cladoxylon
• Some of the
dichotomously
branched, fan-shaped
leaves were fertile.
• Each segment of the
sporangiophore
terminating in a simple
sporangium.
• Homosporous.
58. Sporangia in
Coenopteridales
• Eusporangiate.
• Usually homosporous but Stauroptreis was heterosporous.
• Some had annulus.
Sporangia in Stauropteris
Large, thick –walled sporangia were borne at the terminus of
branches.
S. oldhamia from Upper Carboniferous was homosporous,
but S. Burntislandica from Lower Carboniferous was
heterosporous.
Dehiscence was by means of an apical pore.
Numerous spores.
59. Sporangial & Soral
Characters in Angiopteris
• Fertile leaves are alike the sterile leaves
in form.
• Sporangia are borne in distinct sori on
the abaxial side of the leaves.
• Sori are superficial.
• Oblong rings of free sporangia occur,
each over a vein.
• They are sessile and arise in two rows
beneath veins towards the margins
of the pinnules.
• All the sporangia in a sorus are free
from each other.
(https://tse3.mm.bing.net/th?id=OIP.otyl4nF0hunGfpOQzpqMAgHaE8&pid=Api&P=0&w=253&h=169)
(https://pics.davesgarden.com/pics/2019/07/31/DaylilySLP/cce4f2.
jpg)
60. • Eusporangiate.
• Simple.
• Free sporangia have a band of enlarged cells
of thicker walls.
• Each sporangium has
a several layered thick wall, with stomata in
the outermost layer.
• High spore output: 1450-7000.
• No annulus, sporangia open by slits on the
inner side, or by pores near the apex.
(https://www.fernsoftheworld.com/wp-
content/uploads/2014/09/Angiopteris_video.png)
61. Sporangial Characters in
Ophioglossum
• Sporangia are produced in two rows on
a fertile spike that arises at the junction
of the petiole with the sterile lamina, on
its adaxial surface.
• Spike is a simple, more or less cylindrical
and stalked structure with laterally
fused elongate synangia.
• Generally, a single spike is associated
with a leaf but in O. palmatum several
spikes are present on a single leaf.
• The fertile spike bears two
marginal rows of sunken and laterally
fused sporangia.
(https://ebps.org.uk/wp-content/uploads/2017/06/DSC05036.jpg)
(https://www.delawarewildflowers.org/images/
ophioglossum_vulgatum_2.jpg)
(http://1.bp.blogspot.com/_mFm838BFqBE/TS6qICNB31I/AAAAAAAAAcw/flK9jA1UM5c/s1600/ophioglo
ssum-esporangios8.jpg)
(https://alchetron.com/cdn/ophioglossum-
pusillum-cea403e3-ab09-4eea-a08b-3b793f6ddb2-
resize-750.jpeg)
62. • The apical region of the spike
which is somewhat conical and
devoid of sporangia projects as
a sterile process.
• The length of the spike
varies according to the species.
• Prominent vascular bundle and
stomata are present on spike. A
ring of bundles become
organized within the stalk of the
complex. At higher levels, the
bundles on the adaxial side
become more closely grouped
and enter the fertile spike.
• The number of sporangia in
each spike ranges from 6-20,
according to the size of the
plant.
63. • A number of vascular strands run longitudinally
up the middle and
from these strands develop the lateral branche
s which lead to the sporangia.
• At maturity, each sporangium is spherical
or oval and remains embedded in tissue of the
fertile spike.
• The wall of the sporangium is several layers
thick.
• Stomata are present in the sporangium wall.
• Homosporous.
(https://tse4.mm.bing.net/th?id=OIP.aTaqWvesNGE9gKvx15F
PCAHaFk&pid=Api&P=0&w=236&h=177)
64. • Below it, is tapetum
that comes from the inner
most wall layer.
• No special dehiscence
mechanism. It takes place
by drying out and
shrinking of the sterile
tissue within the spike.
• Each sporangia opens by
a transverse slit in the
sporangial wall.
• Spores as many as
10000.
(https://ebps.org.uk/wp-content/uploads/2017/06/DSC05039.jpg)
65. Morphological Nature of Spike
• Bower (1896) suggested that fertile spike is a single septate
sporangium growing on the adaxial face of the leaf.
• This interpretation was later on (in 1911) rejected by Bower
himself.
• According to present considerations, the fertile spike is
regarded pinna-like in nature.
• It has been held by Goebel (19150, that the spike represents
a single pinna.
• According to Roeper (1859), the spike represents two
basal pinnae whwich have fused laterally, during
the ontogeny to form the spike. The interpretation
is supported by the course of vascular supply to the spike.
The spike exhibits the same vascular supply as would
ordinarily go to a pair of pinnae. Later this interpretation
was supported by Chryslar (1910) and Bower (1926).
• According to Zimmermann (1930), the fertile spike and
sterile lamina are the divisions of a leaf, representing
a dichotomy.
(https://ebps.org.uk/wp-content/uploads/2017/06/DSC05034V2.jpg)
66. Soral & Sporangial
Characters in Osmunda
• Sporangia do not occur in sori and are
attached along the margins of non-laminated
fertile pinnules, forming tassel-like structures:
either on an entire leaf (O. cinnamomea)
on part of it ( O. regalis, O. claytoniana)
• Sporangia are not protected by indusium.
• In O. regalis, the fertile pinnules are found
in the distal regions of the leaf.
• In O. claytoniana, the fertile pinnae are
restricted to the middle of the leaf.
• In O. Cinnamomea, there are two kinds of
leaves, the sterile and fertile. The fertile leaves
are reduced and blade-less.
(https://pics.davesgarden.com/pics/2006/06/03/Equilibrium/7cc
035.jpg)
(https://tse3.mm.bing.net/th?id=OIP.vaYLvrppt0OiThn9QMZ
wrgHaJ4&pid=Api&P=0&w=123&h=164)
(https://www.gardenersdream.co.uk/images/osmunda-regalis-p4672-37141_image.jpg)
67. • Homosporous
• Transitional type of
sporangial development.
• Lack an indusium.
• During the course of sporangium
development, the walls of a group of
jacket cells located near the tip
towards one side of
the jacket become thickened to
form the rudimentary annulus.
• Mature sporangia is a large
pyriform body with short and massive
stalk.
• One-layer thick wall.
(https://i2.wp.com/mgnv.org/wp-
content/uploads/2021/07/Sporangia_Osmunda_spectabilis_fertile_
leaflets_May_MMF.jpg?resize=1536%2C1398&ssl=1)
(https://live.staticflickr.com/6011/5937579433_99ec74f7c5_
b.jpg)
68. • Dehiscence of the sporangium takes
place by vertical split starting from
the rudimentary annulus to the outer
side of the sporangium.
• High spore output: 128-512.
• Spores are spherical with a triradiate
mark and abundant chloroplast.
(https://live.staticflickr.com/2692/4382051069_eae4
a41938_b.jpg)
69. Soral & Sporangial
characters in Marsilea
• Heterosporous.
• Reproductive structures are flattened
bean-shaped, spherical to ovoid,
epipetiolar, stalked sporocarps,
inserted a short distance above the
petiole.
• Length of the stalk and its point
of attachment is variable in
different species.
• Normally there is one sporocarp per
petiole but in forms like M. polycarpa,
there are large number
of sporocarps attached on one side of
petiole and in M. quadrifolia the stalk is
occassionally branched forming 2-
3 sporocarps per petiole.
(https://3.bp.blogspot.com/-gNPth-
RdjmE/WHpYwEldbHI/AAAAAAAAI7I/esevM2uPAZ8ZW14OiyTKMsjVhJkJc_axgCEw/s1600/Marsilea%2Bsporocarp.jpg, https://tse1.explicit.bing.net/th?i
d=OIP.4XSzRMex9vqL2wFM8ZpQugAAAA&pid=Api&P=0&w=246&h=152)
70. • In some species, the sporocarps have distinct external ridge,
the raphe and two bumps.
• The raphe represents the end of attachment of the stalk.
• One of the bumps is stouter and is present at the end of the
raphe.
• The sporocarps have a thick wall resistant to injury and
dessication and are reported to remain viable up to 35-40
years. The outer layer is epidermis made up of single-layered
cuboidal cell with sunken stomata.
• Inside the sporocarp, around the cavity extends a ring
of gelatinous tissue.
• There are rows of elongate sori, on either side,
extending transversely to the long axis of the sporocarp.
• Each sorus is borne on an elongate receptacle which is
attached to the gelatinous ring, at its two ends as well as to
the wall of sporocarp on one side, along its length; the
other side faces the lumen of sporocarp.
• Number of sori in each sporocarp may vary from 2 in M.
aegyptiaca to 20 in M. Quadrifolia.
(https://www.biophotosharing.com/getphoto.php?id=357)
(https://tse3.mm.bing.net/th?id=OIP.2m-
NVRkuOdL8zW0MN4H8nwHaGR&pid=Api&P=0&w=215&h=1
82)
(https://upload.wikimedia.org/wikipedia/commons/thumb/3
/33/Marsilea_expanded_sporocarp.jpg/1024px-
Marsilea_expanded_sporocarp.jpg)
(https://tse1.mm.bing.net/th?id=OIP.86lZ6DX0c8d3W7ycTdQ
SHwHaFj&pid=Api&P=0&w=221&h=166)
71. • Each sorus is enclosed by a
delicate diaphragm, the indusium.
• On the crest of the receptacle are
borne a number
of megasporangia each having
a single large megaspore.
• Along the sides of the
receptacle are borne smaller
microsporangia each
having numerous microspores.
72. •The vascular supply to the sporocarp branches from the
margin of petiolar trace. From the main trace arise
pinnate series of traces, on each side. These lateral bundles
alternate with one another.
•Each of the lateral bundles bifurcates and the vascular supply
to the individual sorus known as placental bundle arises at
the point of forking.
•The placental bundle in turn bifurcates to form dorsal and
ventral traces.
•Towards the lower margin of the sporocarp,
there is a network of veins due to anastomoses of the pinnate
series of bundles.
73. • The sporocarps open only in water,
splitting bivalve-like.
• The process is initiated by the absorption
of water by gelatinous ring which
expands and frees the sori from the wall
of sporocarp by an abscission along the
receptacle. This is followed by
the breaking of ring at one end.
The broken gelatinous ring results into an
elongate worm-like structure dragging
the sori along with it.
• Microspores are small globular cells.
• Megaspores are large, ivory-
white papillate structures enclosed by
gelatinous layer and are visible to naked
eyes.
https://tse4.mm.bing.net/th?id=OIP.MaVu1a-
d_04OPFZsDty6VAAAAA&pid=Api&P=0&w=154&h=154, https://live.staticflickr.com/1383/5120664022_d29d4d5cdf_b.jpg)
74. Morphological Nature of Sporocarp
• Two manin interpretations:
1. Leaf-segment or laminar hypothesis: Sporocarp is a lateral modified
fertile segment of the leaf.
2. Petiolar hypothesis: Sporocarp is homologous with the swollen end
of the petiole of a sterile leaf whose marginal cells differentiate into
sporangial initials instead of the four leaflets.
75. Soral & Sporangial
Characters of Salvinia
• Sporangia are aggregated into sporocarps.
• The sporocarps appear in chains or clusters ranging in number from 4
to 20.
• The sporocarps are usually borne on the inner segments of
submerged leaves.
• In arrangement, the sporocarps are sympodial.
• The shape of the sporocarp varies from globose to ovoid, occasionally
flattened also with ridged surfaces.
• The wall of the sporocarp is made up of two layers of cells. The
outermost layer when young is clad with hairs.
• Externally all sporocarps are alike. But unlike in Marselia, the
sporocarps are mono-sporangiate. (https://tse4.mm.bing.net/th?id=OIP.ZOtkqmEbuAdDKNhn-
jqtOAHaHR&pid=Api&P=0&w=164&h=161, https://tse4.mm.bing.net/th?id=OIP.eFvQgx
vEaqSC1zvQtz7AEwHaFj&pid=Api&P=0&w=240&h=180)
76. • In each sporocarp there is an elevated receptacle bearing
sporangia.
• The receptacle receives a vascular trace from the leaf
segment.
• The first one or two sporocarps in each cluster
are megasporangiate whereas all the later formed ones
are microsporangiate.
• In a megasporangiate sporocarp, only about 25 mega-
sporangia are seen whereas in a microsporangiate
sporocarp, the number is much larger because of the
branching of the receptacle.
• Leptosporangiate.
• Sporangial wall is delicate and single layered, without
annulus.
• As the spores mature, the tapetal cytoplasm encompassing
the spores gets hardened and forms a mass called
‘massula’.
• After the sporangia are mature the sporocarps get detached
from the leaves and sink to the bottom of the pond.
• The sporocarps open up due to the mechanical decay of
their walls. This results in the release of spores, which rise to
the surface where the rest of life cycle is completed.
(https://live.staticflickr.com/7224/7212694442_3f42c67cec_z.jpg, https://pubs.usgs.gov/ds/854/Photograph_collectio
n/Plants/Ferns/Salvinia%20molesta_sporocarps_0008.JPG)
77. Soral & Sporangial
Characters of Azolla
• Heterosporous.
• Sporocarps are dimorphic: mega- & microsporangiate.
• Sporocarps are monosporangiate.
• Sporocarp wall is bilayered and delicate except at the
tip of megasporangiate sporocarp where it is firm.
• Microsporangia are globose, long stalked producing 64
spores.
• Megasporangiate sporocarp contains one
megasporangium which produces 32 spores, all of
which except one degenerates. (https://www.researchgate.net/profile/Yudh-Singh/publication/273169664/figure/fig2/AS:523095527170048@1501727100077/Azolla-microphylla-plants-showing-
sporulation_Q640.jpg, https://www.wnmu.edu/academic/nspages/gilaflora/a_mexicana1.jpg)
78. • In microsporangium, the tapetal plasmodium
develops a number of vacuoles. Each vacuole
becomes hardened and divided by partitions
into number of alveoli. Each hardened
alveolar material within which
the microspores are embedded is called
a massula.
• In some species, the massula develop tubular
outgrowths with the anchor shaped tips
called glochidia. These structures helps in
attaching the microspore massulae to the
megaspore massulae.
• In megasporangium, tapetal plasmodium
divides into 4 massulae. One
of thse envelopes the single megaspore. The
other three massulae are grouped above the
megaspore.
(https://tse1.mm.bing.net/th?id=OIP.hC0fIfz9FNTi_qgZO8dyOAHaGq&pid=Api&P=0&w=197&h=177, https://microscopyofn
ature.com/sites/default/files/2020-01/Kapsel.png)
79. • The presence or
absence, and form of
glochidia on
a massula are important
characters,
delimiting the species.
• The sporangium is
covered by a two
layered inducium.
• Massulae and
megaspores are
released by the
decomposition of
sporocarp wall.
80. Comparing the sporocarps of Marsilea,
Salvinia & Azolla
Marsilea Salvinia Azolla
• Sporocarp is monomorphic • Monomorphic • Dimorphic
• Sporocarp is bisporangiate • Monosporangiate • Monosporangiate
• Heterosporous sporangia • Heterosporous sporangia • Heterosporous sporangia
• Sporocarp wall is 3 layered. • Bilayered • Bilayered
• Thick & resistant wall • Delicate wall, with outer layer
covered with hair.
• Delicate except at the tip
of megasporangiate sporocarp
where it is firm.
• Normally there is one sporocarp
per petiole.
• Sporocarps appear in chains or
clusters ranging in number from 4
to 20
• Sporocarps are developed in
groups of two or four in the axil of
the dorsal lobe of the sporophyll.
• Massulae absent. • As the spores mature, the tapetal
cytoplasm encompassing the
spores gets hardened and forms a
mass called ‘massula’.
• Massula present.
81. • Gloichidia absent • Gloichidia absent. • In some species, the massula develop
tubular outgrowths with the anchor shaped
tips called glochidia. These structures helps
in attaching the microspore massulae to
the megaspore massulae.
• Each sorus is
enclosed by a
delicate
bilayered indusium
.
• Indusium forms the sporocarp
wall.
• Bilayered inducium forms the sporocarp
wall.
• 2-20 sori in each
sporocarp.
• Single sorus in each sporocarp. • Single sorus in each sporocarp.
• Annulus is absent. • Annulus is absent. • Annulus is absent.
• Receptacle is
branched.
• Receptacle in megasporangiate
sporocarp is unbranched whereas
those in microsporangiate
sporocarp is branched.
• In a mega-sporocarp, the mega-sporangium
arises on a small receptacle at the base. In a
micro-sporocarp there is a central cushion
like receptacle which gives rise to a number
of microsporangia.
82. Soral & Sporangial
Characters in Pteris
• Homosporous.
• The sorus of Pteris is called coenosorus .
• Coenosori are marginal, borne continuously on a
vascular commissure connected with vein ends.
• The sporangia of Pteris form a continuous linear
sorus along the margin, hence the individuality of
sori is lost.
• The coenosori are protected by the reflexed margin
(false indusium) of the pinnae.
• Sori are of mixed type intermingled with many
sterile hairs in between the sporangia.
• Leptosporangiate type.
• A mature sporangium has a long stalk that
terminates in a capsule
(https://tse3.mm.bing.net/th?id=OIP.7q6bHUIh8LWSMuxIyuF8SAHaFj&pid=Api&P=0&w=217&h=163, https://fernsofmacaronesia.org.
uk/wp-content/uploads/2020/06/Pteris-vittata-6-008-1050x700.jpg)
83. • The jacket of the capsule is single-layered, but with
three different types of cells:
(1) A thick walled vertical annulus incompletely
overarches the sporangium
(2) A thin-walled radially arranged stomium
(3) Large parenchymatous cells with undulated walls.
• Spores are triangular in shape with trilete aperture,
bounded by two walls. The outer wall, exine, is
variously ornamented.
• The sporangium dehisces transversely along the
stomium due to the shrinkage of annular cells.
• Each sporangia produces 48 spores.
84. Soral & Sporangial Characters
in Lygodium
• Homosporous.
• Sporangia which are produced in abundance
are borne in two rows along the ultimate
veins.
• They form spike like structures on the leaf
margin.
• A single sporangium occur at the end of
veinlet and each is covered by a flap.
• The flap is a false indusium representing an
extension of leaf margin.
• The sporangia are born singly, forming
“monosporangial sori”
(https://tse2.mm.bing.net/th?id=OIP.b6OAas7gaR7QfY2fMVC1UAHaHa&pid=Api&P=0&w=157&h=157, https://tse2.mm.bing.net/th?id=OIP.7EtS0E8owimX-
gjIImlrqgAAAA&pid=Api&P=0&w=192&h=152, https://tse3.mm.bing.net/th?id=OIP.hHJXqpfPnZWZrfb0NNWMagAAAA&pid=Api&P=0&w=230&h=185)
85. • Each sporangium has a short and thick stalk
with a spore number of 128 –256.
• Leptosporangiate development
• Massive bilayered tapetum.
• The annulus is a terminal group of thick –
walled cells, confined to distal end
of sporangia.
• The dehiscence is longitudinal , most
primitive type in ferns.
• Capsule of the sporangium is pyriform and
curved.
• Definite stomium is present and the
dehiscence is longitudinal.
• Spores are usually triangular, tetrahedral and
trilete.
• Exine is striate.
86. Soral & Sporangial Characters
in Acrostichum
• The upper pinnae are small and fertile, whereas
the lower pinnae are infertile.
• The length of the lowest pinna is occasionally
smaller than that of the upper pinnae.
• Sporangia cover the lower surface of the fertile
pinnae or are located at the tip.
• The sporangia have many sterile, multicellular
filaments that are threadlike, with darkened cells
at the end.
• This genus gave rise to the word acrostichoid, due
to the tightly packed sporangia on the abaxial
side of the lamina (Tindale, 2009).
• Acrosticoid sorus is the most advanced sorus.
https://tse3.mm.bing.net/th?id=OIP.aRs20K90A9nl0_0UEVYjEgHaE8&pid=Api&P=0&w=236&h=157, http://www.discoverlife.org/IM/I_GY
/0007/320/Acrostichum_danaeifolium,_sori,I_GY741.jpg, https://tse2.mm.bing.net/th?id=OIP.X61wz3E-
gdP2kLo_wDk0nQHaJ4&pid=Api&P=0&w=133&h=178)
87. • The presence of sporangia and
the absence of
sori differentiate Acrostichum fro
m other ferns (Hill, 2006).
• A mature sporangium has a long
stalk that terminates in a capsule.
• Exindusiate.
• Mixed type of sori.
• Spores yellow, tetrahedral,
minutely tuberculate or roughene
d, equatorial flange absent.
• Spores clear to translucent, 1 –
1.5 mm wide, and buoyant.
(https://biogeodb.stri.si.edu/bioinformatics/dfmfiles/files/c/25134/25134.jpg)
88. Soral & Sporangial Characters in
Gleichenia
• Sporangia occur in ring-shaped sori on the abaxial surface of the
pinnule.
• Ex-indusiate.
• No segregation of sterile and fertile leaf parts.
• Sori are naked and lie in two rows, one on either side of the mid-
rib of the pinnule.
• Number of sori on a pinnule varies from 1 to many.
• Sporangia arise on somewhat elevated circular receptacle and
vary in number from 4 to 15.
• Size of the sporangia varies with their number within a sorus.
• In the species which have few sporangia, the sporangia occur in
a uniseriate ring at the margin and the centre of the sorus is
vacant. (https://live.staticflickr.com/3450/3186416993_1548b1864e_b.jpg, https://blog.tepapa.govt.nz/wp-
content/uploads/2013/02/glechenia_inclusisora_6374_sori_0011.jpg, https://live.staticflickr.com/3810/10328706023_0dc4b4f66a.jpg, http://www.nzflora.info/factsheet/images/Image1YX8large.jpg)
89. • In other species, the sporangia found in the centre, lying on top
of others.
• When examined superficially, a sorus looks like a synangium, but
it is not so.
• Mature sporangium is pyriform.
• Young sporangia are usually covered by hairs or scales.
• Sporangia lie on two sides of the midrib of the pinnules.
• Some cells of the jacket are thick-walled and form a distinct
annulus whose position is oblique horizontal.
• Simple sori.
• Leptosporangiate development.
• Short and stout stalked sporangia.
• Distinct annulus present.
(https://www.yarraranges.vic.gov.au/files/assets/plants/ferns/605b-2-gleichenia-microphylla-mb.jpg))
90. • Single-layered jacket enclosing varying
number of spores.
• Spore outcome:128-1024.
• Each sporangium has a large number of
spores and has an obliquely transverse
complete annulus which causes
longitudinal dehiscence.
• Spores are trilete in
G. circinata whereas monolete in
G. bifidus.
• Homosporous.
• Has slightly papillate surface.
• Spores vary in size according to the
species.
(http://www.nzflora.info/factsheet/images/Image1YX7large.jpg)
91. Soral & Sporangial
Characters in Adiantum
• There is no distinction into fertile and
sterile leaves.
• Soru bearing leaves are called
sporophylls.
• The sori are born superficially at the
distal end of the pinnae. But the sori
are not exactly marginal. They are
borne a little behind the tip of the
veins, submarginal.
• The sorus bearing margin of the leaf
incurls and forms the false inducium.
• The sorus is of the mixed type.
(https://floracanaria.com/fotos/Adiantaceae/Adiantum_renif
orme/image/Adiantum_reniforme_08.jpg)
(https://live.staticflickr.com/3477/3809777711_f1bd181c24_b.jpg)
(http://1.bp.blogspot.com/-
dkVkvx3CJE0/TbCriOkZD1I/AAAAAAAAAyo/4f2RalVdzA4/s1600/hispi
dulum-soros.jpg)
92. • A
mature sporangiuim has
a stalk made up of three
rows of cells.
• The stalk terminates in
a globose or biconvex
capsule.
• The wall is single
layered.
• The sori are covered
and protected by reflexed
margins of leaflets, which
are membranous
structures called false
indusium.
(https://cdn.auckland.ac.nz/assets/nzplants/Ferns/Some%20Native%20Ferns/Pteridaceae/Adiantum%20cu
nninghamii/9-%20adiantum_cunn_lflet5(PS7).jpg)
93. • Each sporangium is oblong and consists of
a multicellular stalk and the head or
capsule.
• Capsule is biconvex.
• Distinct vertical annulus.
• The annulus is separated from the stalk by
two or three cells.
• The stomium is separated from both the
stalk, and the annulus.
• The rest of the sporangial wall is
composed of a few large cells.
• There is an obliquely vertical annulus of
12-24 cells long.
• The sporangium dehisces transversely
liberating the spores.
• Leptosporangiate development.
• Homosporous.
• Spores are tetrahedral in shape with a
triradiate ridge and spore wall is two
layered
(https://tse4.mm.bing.net/th?id=OIP.ggTtW11L00QFgi9S1AUx2QHaE9&pid=Api&P=0&w=237&h=159)
95. Reference
1. Yatskievych, George. (2003). Pteridophytes (Ferns).
10.1038/npg.els.0003679.
2. Vashishta, P. C. .Botany For Degree Students Pteridophyta. New
Delhi: S. Chand & Company Ltd.,2002.
3. Pandey, S. N, Misra, S. P & Trivedi, P. S. A Textbook of Botany.
Thirteenth Edition. Noida, India: Vikas Publishing House Pvt. Ltd.,Vol
2,Part 2, 2016.
4. Rashid, A. An Introduction to Pteridophyta. Second Revised
Edition. New Delhi: Vikas Publishing House Pvt. Ltd., 1999.