NonVascular and Seedless Vascular Plants

ADAPTATIONS TO LIFE ON LAND Making the transition from a small, aquatic, Chlorophyta alga to the large, multicelled terrestrial plants of today required a number of structural and physiological changes as outlined below. Recent* findings support the view that the most ancestral green flagellate was biflagellate and asymmetric, and possessed an eyespot, square scales and multilayered structures. *Lemieux, C., Otis, C.,and Turmel, M. Nature 403,649-652 (2000). This led to a number of evolutionary dead ends possessing many of the characteristics present in the higher plants. Conditions of Land Life To maintain itself a living cell must obtain water and other materials by absorption through the cell membrane. Materials must be available fairly consistently throughout all cells of an active organism. Diffusion can be used to distribute materials if the organism is small or consists of only a few cells. Difficulties arise in larger organisms because some cells are internal - out of direct contact with the environment. These problems are aggravated when some parts extend into the air and are distant from a source of water and dissolved materials. Water must be passed along from cell to cell by osmosis and diffusion. These are short distance processes which are slow and ineffective over distances inside the plant. Movement of materials across great distances is the first barrier to any plant attempting to extend itself into the air. Development of vascular systems somewhat analogous to circulatory systems in animals has overcome this problem in higher or vascular plants. Specialization at the level of the tissues and organs was also necessary. Systems to gather in nutrients and water (roots) and collection and conversion systems to utilize the power of sunlight (leaves)and systems to hold the plant upright and increase its competitive ability to battle for sunlight (stem) were required. Changes in reproductive structures and behaviors led to protected male gametophytes (pollen) and embryos that were protected and nurtured (seeds). A COMMON SET OF FEATURES EVOLVED BY GREEN PLANTS IN RESPONSE TO THE LAND ENVIRONMENT 1. Cuticle - waterproofing of outer layers of leaf and stem 2. Pores or Stomates surrounded by Guard cells and regulated by K+ levels - aided in water balance within leaf and entire plant body through transpiration 3. Rhizoids or roots - provided anchorage, absorption, support 4. Lignin - strengthening between cells and within tissues 5. Vascular tissue - xylem and phloem; move water and nutrient upwards and food downward 6. Resistant spores and seeds - prevent dessication, contain inhibitory substances; ensure survival of the species 7. Gamete forming cells enclosed within a jacket of sterile cells provides protection from water loss 8. Embryos formed and held within parent tissues - provides protection and a food supply to developing plant 9. Root systems show a highly refined geotropic response. EVOLUTIONARY TRENDS ACCOMPANYING THE TRANSITION

1. The sporophyte has replaced the gametophyte as the dominant stage in the life history of the plant. During this shift, the size of the sporophyte progressively increased, while that of the gametophyte decreased. 2. The increase in the size of the sporophyte was accompanied by an increase in strength and efficiency of its vascular tissue. 3. As tissues became more specialized, new organs were added progressively: simple bodies with rhizoids ---> stems ---> stems with roots and tiny leaves ---> manyveined leaves ---> woody tissues 4. The male gametophyte evolved into a waterproof pollen grain, which traveled to the female gametophyte before releasing the sperm. Sperm no longer faced a long, dangerous swim to the egg. 5. The female gametophyte of all land plants retains the egg and protects the zygote as it develops into an embryo; in higher land plants, the female gametophyte itself is retained on or in the sporophyte parent, which contributes food and protective coatings to the seed, a new dispersal structure containing the embryo of the next sporophyte generation. 6. In the higher vascular plants, the spore, a single haploid reproductive cell that is the dispersal stage of lower land plants, has become differentiated into megaspores, which give rise to female gametophytes, and microspores which give rise to pollen grains (male gametophytes). Instead of being shed, the megaspores remain in the sporophyte, which protects the female gametophyte and plays a major role in the production of the seed. Seeds replaced spores as the dispersal stage in the life history. 7. As plant structure and reproduction became more and more independent of water and moisture in the environment, plants underwent adaptive radiation and spread into more and more different habitats. Sexual reproduction (fusion of GAMETES) and Asexual reproduction (by means of SPORES), occur in ALTERNATING GENERATIONS. One generation, where the plant body is known as the GAMETOPHYTE is Haploid (n) and produces Haploid GAMETES, the Sperm and the Egg. These gametes then fuse to form a ZYGOTE in the process of FERTILIZATION. This Diploid Zygote develops into a SPOROPHYTE (also Diploid). The Sporophyte undergoes MEIOSIS and forms SPORES that develop into an adult organism called the GAMETOPHYTE This completes the cycle. The zygote was formed as a result of the fusion of two cells, while the Gametophyte plant developed as the result of asexual reproduction (mitosis) of the spore. METAPHYTA: NON-VASCULAR PLANTS - THE BRYOPHYTES All available evidence seems to indicate that this division represents an evolutionary dead end. There is no evidence to indicate that this division is ancestral to any of the modern vascular plants. It has been suggested that differences between the three major classes within the division may necessitate reclassification into separate divisions at some time in the future. These plants show many of the adaptations needed for a successful transition to a terrestrial environment. As a division these plants: a. lack specialized vascular tissue or supportive tissue

b. possess a characteristic life cycle which follows the generalized life cycle where the SPOROPHYTE is attached to and dependent upon the GAMETOPHYTE for nutrition - the GAMETOPHYTE is dominant, conspicuous, and longer lasting In higher, vascular plants the relative importance of the SPOROPHYTE and GAMETOPHYTE are reversed. Gametophyte plants may be MONOECIUS (one house) and contain both male and female organs or, DIOECIOUS (two houses) have separate plants for each. The sex organs are multicellular and bounded by a layer of sterile cells. Bryophytes are usually small, ranging from 2-20 cm. They lack true roots, stems, or leaves although they have structures which accomplish many of the functions of these organs. Leaf-like structures are usually a single cell layer thick and are called PHYLLIDS. Root-like structures, called RHIZOIDS, serve mainly for anchorage, and stems or stalks, called SETA or CAULIDS, serve for support. There are three recognized classes of Bryophyta: Liverworts, Hornworts, and Mosses Class Musci - The True Mosses This class is frequently confused with sea moss (red algae), reindeer moss (lichen), spanish moss (flowering plant), and club mosses (primitive vascular plant). Mosses are distinguished from Liverworts and Hornworts by: 1. an algal-like PROTONEMA 2. radial symmetry of the plant body 3. the elaborate capsule of the mature sporophyte There are approximately 670 genera representing about 16,000 species of mosses. They are normally found in wooded, moist areas, usually in dense stands forming carpets or mats, and are seldom more than 6-8 inches tall. There are some species found in desert areas. They are sensitive to air pollution, especially sulfur dioxide. Economically they are of importance as: uccession

Sphagnum moss, which has a high water holding capacity and adds some acidity to the soil The following characteristics of mosses distinguish them as transition plants between aquatic and terrestrial habitats. 1. dependence on water for sexual reproduction

2. vegetatively well adapted for life on land 3. relative size and independence of sporophyte and gametophyte generations; situation reversed in higher plants It appears that the sporophyte, which eventually evolved a vascular system in higher plants, is better adapted to life on land (terrestrial), in that spore production is well suited to that type of habitat. By contrast, the biflagellated gametes of the gametophyte generation are better suited to an aquatic mode of life. Mosses seem to retain a close algal relationship as evidenced by the structure of the PROTONEMA, similar to that of some green algae; possible evidence for evolution from green algal forms. Mosses established at least two critical features which later were more fully developed by land vascular plants. 1. retention and protection of the embryo within the female parent plant 2. presence of a strand of tissue occupying the center of the stemlike axis used for support and conduction - may be prevascular tissue which evolved into true vascular tissue containing tracheids and sieve tubes; in mosses this central strand of tissue consists of cells called HYROIDS ALTERNATION OF GENERATIONS in Mosses Haploid spores form within the capsule which is covered by a lid called an OPERCULUM. As many as 50 million spores may be formed. When released the spores develop into a branched, filamentous PROTONEMA from which a leafy gametophyte develops (may be monoecious or dioecious). Biflagellated sperm are released from the ANTHERIDIUM and attracted to the ARCHEGONIUM by malic acid. Within the archegonium the sperm fertilizes the egg cell which develops into a zygote which divides mitotically to form the SPOROPHYTE. Typical structures form and spores develop within the capsule. Some species have a PERISTOME beneath the operculum (humidity sensitive hairs regulating spore dispersal (hygroscopic), e.g. the genus Polytrichum. METAPHYTA: SEEDLESS VASCULAR PLANTS Tracheophytes (vascular plants) are the largest group of plants on earth and are characterized by the presence of tracheary elements and sieve tube elements organized into xylem and phloem in the adult sporophyte. (see notes above on Plant Adaptations to the Land Environment) All lines of evidence suggest that the tracheophytes developed directly from some complex, multicellular Chlorophyta. Unfortunately, an extremely large gap exists between the algae and vascular plants, and there are no living plants which bridge the gap. Bryophytes are somewhat intermediate but are closer to algae than even the simplest tracheophytes, and it appears that the intermediate plants have become extinct. In addition, tracheophytes are identified by independent and dominant sporophytes and greatly reduced gametophytes. This is in direct contrast to the Bryophytes. The sporophyte of most tracheophytes are differentiated into roots, stems, and leaves. Also, vascular plants are typically land plants and, like bryophytes, have evolved terrestrial adaptations. Check out this comparative

information on the lower tracheophytes. Evolution in the Lower Tracheophytes (Psilophytes, Lycophytes, Sphenophytes, and Ferns) Transition from water to land was gradual over millions of years. Those plants which evolved adaptations permitting growth in a dry environment established themselves as the first land plants. The basic problem confronting early land plants was availability of water. The tracheophytes did not have the limitless supply of water which was available to the aquatic green algae. A successful land plant must possess an efficient mechanism for absorbing materials from the soil and transporting them throughout the plant. Once obtained from the environment the water must be conserved. Division Psilophyta The evolutionary significance of the Psilophytes is that they are the most primitive known group of vascular plants and are considered to be a transitional group linking the aquatic algae with more advanced tracheophytes (club mosses, horsetails, and ferns). In some respects, the Psilophytes substantiate the hypothesis for the evolution of the root as proposed by Lignier. In other respects they are of interest because they suggest how club mosses, horsetails, and ferns may have originated by various leaf modifications and developments. Certain psilophytes with small leaves developed as emergences and may have given rise to the Lycophyta line. Others with whorled branches may have been the forerunners of the Sphenophyta line. The fern line may have developed from forms in which the branch tips were flattened, possibly leading to the evolution of large leaves. Two major genera represent this group, Psilotum and Tmesipteris. Division Lycophyta Represented by Lycopodium and Selaginella,the lycophytes are believed to have evolved from the psilophytes. Although members of this group are of no real economic import, the group does illustrate some noteworthy evolutionary advances over the psilophytes. The presence of true roots, and leaves, increased devel opment of the vascular tissue, and the organization of sporophylls into cones (strobili) are all viewed as advances over the psilophytes. It is not clear why the lycophytes have not given rise to more advanced groups of plants. Some have developed secondary growth and exhibit early stages of the seed habit. Division Sphenophyta These plants are of little economic significance, although together with the lycophytes, they contributed their vegetative parts to the formation of coal during the Carboniferous period. They are considered to be a separate line derived from the psilophytes which did not give rise to other plant groups. Major genus is Equisetum, known as horestails or scouring rushes. Some species accumulate silicaceous materials in their cell walls. Division Pterophyta (Ferns) Ferns and other pterophytes are believed to have developed from the psilophytes. Certain fossil ferns resemble the psilophytes rather closely. The seed plants are considered to have derived from certain of the extinct fern class. Modern fern groups are of little economic importance. From an aesthetic view, they are grown exclusively for decorations. The fiddleheads and young foliage of

many species are edible and are used as green vegetables in the Orient. A drug is derived from the rhizomes (underground stem) of some ferns and is used to expel worms (vermifuge) especially tapeworms, from the intestinal tract of man. Fossil ferns contributed to the deposits of the Carboniferous period.

part is surrounded byan upgrowth of the thallus,the involucre. The development of the archesporium beginsas a dome-like layer within the summit of thesporophyte. The sporophyte continues togrow forseveral weeks or longer from a meristem close toits base. During this growththe archesporium differentiates basipetally as a hollow cylinder, thecenter of which is occupied byasterile columella.Multicellular elaters (which lack distinct spiralthickenings and are often referred toas pseudo-elaters) differentiate amongst the sporogenouscells. When the spores at the top of the capsule areripe, the capsule dehisces basipetally (Fig. 5.15a)along two longitudinal slits, the opening begin-ning near the tip. As the upper part of the capsuledries, the valves separate completely above, andtheybegin totwist longitudinally.The consequentcontortions of the valves expose the spores andelaters adhering tothe central column, and dis-persal begins. The separation of the valves contin-ues downward as the spores mature, and mean- while the basal meristem generates newsporophytic tissue at about the same rate.Consequently a single sporophyte continues to yield spores over a considerable period. The sporeshaveconspicuously thickened and sculptured walls. Anthoceros and its allies further differ fromother liverworts, but resemble the mosses, in pos-sessing photosynthetic tissue in the outer layersof the extending sporophyte. Stomata are alsopresent, as in the capsules of some mosses. Thesporophyte is thus not entirely dependent on thegametophyte for nutrition, but, since the sporo-phyte will still mature even if it is covered with atinfoil cap, it seems likely that it is able todraw asubstantial proportion of its essential metabolitesfrom the parent gametophyte. e v o l u t i o n a r y p o s i t i o n The Anthocerotales are remarkable amongst theliverworts in recalling the features of both thealgae (the presence of the pyrenoid in the chloro-plast) and the mosses and higher plants (the pres116THE SUBKINGDOM EMBRYOPHYTA: DIVISION BRYOPHYTA (MOSSES AND LIVERWORTS) Figure 5.15. Anthoceros laevis . (a) Female thallus withsporophyte. (b) Section of antheridial chamber. (c) Section of nearly mature archegonium. (d) Diagrammatic longitudinalsection of a sporophyte showing the different regions.

ence of stomata and the continued growth of thesporophyte). They are consequently thought bys o m e t o s t a n d c l o s e t o t h e l i n e o f e v o l u t i o n leading from the algae toterrestrial vegetation. The intermediate position of the Anthocerotalesextends even totheir ultrastructure. The electronmicroscope conrms that the chloroplasts of Anthoceros resemble those of the algae, but thechloroplasts of Megaceros , which are several ineach cell, often lack pyrenoids and possess well-dened grana. They thus resemble those of landplants generally. Another feature of the Anthocerotales whichhas excited much attention is the axial formof the sporophyte, and the tendency in some speciesfor the foot toproduce rhizoidlike outgrowthsinto the tissue of the gametophyte. Since thesporophyte in some forms is long-lived and mayeven persist for a time after the death of theparent gametophyte, it may indicate how simpleaxial plants, such as those of the Silurian andearly Devonian (p. 135), haveevolved. Alternativelythe Anthoceros condition may be derived, thesporophyte having become reduced and almostdeprived of its independence. Although rm evidence relating to either pos-sibility is lacking, it seems quite plausible that ag r o w t h f o r m s i m i l a r t o t h a t s h o w n b y t h e l i v i n g Anthocerotales did play some part in the evolu-tion of land plants from algal ancestors.Unfortunately the only fossils so far known attributable tothe Anthocerotales are spores from the Tertiary. Bryopsida (mosses) The mosses are a class much greater in numberand more widely distributed than the liverworts,occurring in almost every habitat supporting life. Apart from being the dominant vegetation in acid bogs, and alpine and arctic regions, theyare afamiliar feature of woodlands and hedgerows. B R Y O P S I D A ( M O S S E S ) 1 1 7 Figure 5.16. Anthoceros laevis . Female plants bearing youngsporophytes. Scale bar 5 cm.

Some species even survive the polluted atmos-phere of urban areas, often forming dark greencushions between paving stones and in otherdamp crevices. Mosses in exposed situations,including Sphagnum in bogs, often reach tempera-tures of 3040C (86104F). Isoprene production(p. 11), which is more common in mosses than inany other major group of land plants, increases inthese conditions, and may consume up to8percent of the carbon xed. As in other landplants, isoprene appears to be concerned withprotecting the photosynthetic apparatus fromexcessive insolation and heating. The features which distinguish the mossesfrom the liverworts are found in both gameto-phyte and sporophyte. The protonemal stage of the gametophyte is often conspicuous, the matureformof the gametophyte is always leafy, and therhizoids are multicellular. The sporophyte growsfrom an apical cell. The capsule in many instanceshas a complex opening mechanism which affectsthe distribution of the spores, and sterile elatersare never present. The three orders, Sphagnales, Andreaeales andBryales, differ principally in the nature of theprotonema and the structure of the capsule. There are also differences in the ultrastructuraldetail of the spermatozoids (which resemblethose of other bryophytes) which may havetaxo-nomic signicance. The Sphagnales The Sphagnales, a verydistinctive order, are repre-s e n t e d b y a s i n g l e g e n u s , Sphagnum , conned toacid, waterlogged habitats. The more or less con-tinuous spongy layer of peat bogs consists verylargely of a range of Sphagnum species. The adult gametophyte comprises an uprightmain axis from which whorls of branches arise atregular intervals (Fig. 5.17). The leaves, which are 118THE SUBKINGDOM EMBRYOPHYTA: DIVISION BRYOPHYTA (MOSSES AND LIVERWORTS) Figure 5.17. Sphagnum mbriatum . (a) Upper portion of shoot with sporophytes. (b) Stem leaf. (c) Branch leaf. (d)Arrangement of cells in leaf, adaxial surface. (e) Section of leaf.

closely inserted, haveapeculiar structure whichis diagnostic of the genus, and also, in its nerdetails, of the many species (Fig. 5.17c and d). When rstformed, the leaves are made up of many diamond-shaped cells. These then cut off narrow daughter cells, but on two sides only.Thedaughter cells develop chloroplasts, but themother cell remains colorless. It eventually dies, but before doing so pores are formed in its upperand lower walls and spiral bands of thickening areoften laid down around the cell. The architectureof the leaf of Sphagnum thus provides a splendidexample of the phenomenon of apoptosis , apattern of programmed cell death being imposedupon the developing leaf. The cell-to-cell inu-ences which determine this pattern are as yet wholly unknown. The epidermis of the branches takes the formof a cylindrical sheet of empty elongated cells(termed retort cells), each with an apical poreopen tothe exterior. There is no evidence thatthese cells are formed byasequence of divisionssimilar tothat seen in the development of theempty cells of the leaf. The empty cells of the mature leaf and branches are able totake up water and act as res-ervoirs. The weight of a saturated plant may be asmuch as 20 times that of the same plant dry.Thisunique water-storing capacity of the leavesaccounts for the outstanding bog-building proper-ties of Sphagnum . The acidity of the bogs is largelyattributable tothe presence of phenolic sub-stances in the walls of the dead cells which act inthe same way as ion-exchange resins. The growingplants extract metallic cations from inowing waters and release hydrogen ions, so maintainingthe acidity at values of around pH 4.0 or less. Thelack of oxygen in the lower levels of the bogimpedes oxidation of the organic matter, leadingto accumulations of peat of considerable depth. Well-preserved leaves similar tothose of Sphagnum havebeen found in Permian deposits inRussia, indicating beyond doubt the great antiq-uity of this kind of construction. r e p r o d u c t i o n The reproduction of Sphagnum is probably princi-pally vegetative, the decay of the older parts even-tually causing branches toseparate and thus to become new individuals. Disks of Sphagnum peatfrom depths as great as 30cm (12in.) will produceinnovative growths when exposed tolight and asaturated atmosphere. These probably arise fromthe outer cortex of the buried stems. Some cellsevidently remain viable and capable of division,possibly for as long as 60 years.Mature plants in favorable situations producesex organs freely.Both monoecious and dioeciousspecies occur. The antheridia, each of which begins its development from a single apical cell,lie in the axils of leaves towardthe tips of smallupper branches (Fig. 5.18). These antheridial branches are often stronglypigmented and clus-tered in a conspicuous comal tuft. The femaleinorescence consists of a bud-like aggregateof archegonia and bracts borne laterally near thesummit of the main stem. After fertilization the zygote yields a sporo-phyte (Fig. 5.17a) consisting principally of acapsule, containing a dome-shaped archespor-ium, and a foot. The seta remains inconspicuous,and the function of elevating the capsule is takenon by the base of the female inorescence which,as the capsule matures, grows up as a leaess axis,or pseudopodium (Fig. 5.19). The absence of any cellextension in the seta, and correspondingly of anymarked ux of uid into the developing sporo-phyte, is correlated with the absence of labyrin-thine walls at the interface of the two generationsin the foot.Release of the spores is brought about byairpressure which builds up in the lower half of thecapsule as it dries (and may reach a level of 300kPa(3atm)). Eventually this

pressure is sufcient todislodge the clearly differentiated lid (operculum) with explosive force, and the spores are effectivelydispersed.Developing Sphagnum capsules are occasion-a l l y p a r a s i t i z e d b y a f u n g u s ( Bryophytomyces ). Although the mature capsule may appear perfect,the small spores of the fungus replace those of thehost. This gaverise toreports of heterospory in Sphagnum before the situation was understood. Sphagnum s p o r e s g e r m i n a t e t o f o r m a lament, but this is rapidly replaced b y a s m a l l thallose protonema. This in turn gives rise toa bud which develops into the familiar leafy gamet-ophyte, the protonema meanwhile becoming BRYOPSIDA (MOSSES) 119

moribund and disappearing. Protonemata alsoappear on disks of old peat maintained in humidconditions. Many appear toarise from spores within the compacted peat which haveremained viable. Others are secondary in origin from viablecells in the old stems. The Andreaeales The Andreaeales are an order containing onlythree genera, distinguished bytheir peculiar cap-sules. The leafy gametophyte of Andreaea (Fig. 5.20)rarelyexceeds 1cm (0.4in.) in height. It is usuallyfound growing on rock, chiey in cold, exposedand relatively dry regions. The leaves are olive- brown in color, composed of rounded cells, and inmost species showing no distinct midrib.Sex organs are formed apically.The sporophyteresembles that of Sphagnum in having a domedarchesporium (Fig. 5.20c), and in being borne on apseudopodium at maturity. Dehiscence of thecapsule takes place byfour longitudinal slits which do not meet at the tip (Fig. 5.20b). Thehygroscopic properties of the wall cause the slitsto close in damp conditions, and to open again indry (Fig. 5.20a). The protonema of Andreaea is similar tothat of Sphagnum . Also placed in the Andreaeales is the recently 120THE SUBKINGDOM EMBRYOPHYTA: DIVISION BRYOPHYTA (MOSSES AND LIVERWORTS) Figure 5.18. Sphagnum sp. (a) Antheridial branch. (b)Longitudinal section of antheridial branch. (c) Longitudinalsection of archegonial branch. Figure 5.19. Sphagnum sp. Longitudinal section of nearlymature sporophyte.

discovered Andreaeobryum , growing in the oceanicnorthwestof Canada and Alaska. The capsule iselevated byaseta, and opens less regularly than in Andreaea . The protonema produces curious cylin-drical multicellular processes which are greenand presumably photosynthetic. Whereas Andreaea is usually found on siliceous rocks, Andreaeobryum prefers a calcareous habitat.Current evidence points to Takakia (p. 115) also being correctly placed here. This has a smallupright stem, rising from a liform rhizome,growing on soil partly covering rocks. It isconned to subalpine situations in Japan and theoceanic northwestof the American continent. The stems bear linear appendages (phyllids),usually in groups of two or three. The archegoniaare terminal. The archesporium is domed, and thecapsule lacksaperistome and lid. Dehiscence of t h e c a p s u l e t a k e s p l a c e b y a s i n g l e s l i t w h i c h extends spirally from the base of the capsule tothe tip, a feature unique in the bryophytes. Afterrupture, the calyptra is carried up at the summitof the capsule, but becomes loose and fragmen-tary. The seta remains short and thick and doesnot attain the delicacy seen in many bryaleanmosses. In structure and shape the foot resemblesthat of Andreaea . Although the recognition of Takakia as a moss appears well founded, a distantafnity with the Calobryales, particularly inrespect of the archegonia terminating the leafyaxes, remains a possibility. The Bryales The 600 or so genera of the Bryales formawell-dened order. Although there is a common basicmorphology and life cycle within the order, the variation in size, detailed structure and habitatpreferences is considerable. Many mosses areconned topermanently damp situations in woodlands and bysprings, but others, for example Tortula ruraliformis , are able to survive periods of drought in sand dunes and other arid habitats. The cells of these species appear tohave acquiredthe capacity tocontinue metabolism at a reducedr a t e w h i l e p a r t i a l l y dehydrated. Atthe other B R Y O P S I D A ( M O S S E S ) 1 2 1 Figure 5.20. Andreaea nivalis . (a) Habit of fertile plantshowing dehisced capsule in dry condition. (b) Dehiscedcapsule in wet condition. (c) Longitudinal section of maturesporophyte.