Stelar System in Pteridophytes The central vascular cylinder of the primary axis of pteridophytes is usually referred to as stele (Greek stele = column). Besides xylem and phloem, it includes pith (if present) and is delimited from the cortex by the pericycle, The concept that stele is the fundamental unit of vascular system was put forward by van Tieghem and Douliot (1886). They proposed the stelar theory, according to which the root and stem have the same basic structure consisting of two fundamental units — the cortex and the central cylinder. Van Tieghem considered endodermis (the innermost layer of the cortex) to be the anatomical boundary between these two fundamental units, Although 4 distinct endodermis is present between the cortex and stele in vascular cryptogams, higher plants usually lack a distinct endodermis and pericycle, Esau (1953) and Foster and Gifford (1963), therefore, opined that in relation to higher plants, the term vascular cylinder is better suited than stele. Ih lower vascular cryptogams like Lycopodium and Selaginella, the stele of the axis and the leaf traces develop independently, the stele being mainly cauline and the leaf traces are attached only to its Surface. Contrary to this, the stele of higher vascular cryptogams (8. Preridium, Pteris) is a composite structure consisting of the “ascular cylinder of the stem and leaf traces. TYPES OF STELES IN PTERIDOPHYTES Schmidt (1982) recognized the two principal types of steles in Pleridophytes —(1) protostele, (2) siphonostele.238 Stelar System in Preri —_ —. ——————————————[_“ is Protostele It is a non-medullated stele consisting of a central core of xylem, surrounded by a band of phloem. There is a single or multiple layer of pericycle outside the phloem which is delimited externally by acontinuous sheath of endodermis, The protostele is considered to be the most primitive, both phylogenetically as well as ontogenetically. Fossil Psilophytales (e.g., Rhynia, Horneophyton), as well as many living primitive vascular _plants (e.g., Psilotum, Tmesipteris), characteristically show this type of stele. In Filicales (which are otherwise advanced vascular cryptogams), protostele is seen generally in the sporelings, but is also permanently maintained in the adult stems of some relatively primitive Filicales. The protostele gives off leaf traces without any break in the continuity of the enveloping endodermis. With the departure of leaf trace there is no marked change in the solid xylem core. cortex Sear, h leaf trace endodermi The following four types of protos recognized in pteridophytes. surrounded by its 4, vn teles are [A] Haplostele It is the simplest and most primitive protostele It consists of a solid xylem core smooth circular outline, which is surrounded ring of phloem (Fig. 1A). Haplostele is founda fossil (e-g., Rhynia, Horneophyton, Cooksona) « well as many living pteridophytes (2. Psiotog Selaginella, Lycopodium). [B] Actinostele In this type of protostele the xylem is star-shaped with many radiating arms. The phloem, instead of forming a continuous ring, is present in sma] patches in between the radiating arms of the xylem (Fig. 1B). Actinostele is characteristic of many leaf traces Seti [ Fig. 1 A-D. Different types of protosteles ; A. Hay D. At C, Plectostele, D, Mixed protostele, ee aEa, Fig 2 Transverse section of stem of Osmunda regalis, sowing mixed pith (adopted from Bower, 1923). living (e.g., Psilotum, Lycopodium serratum) and fossil forms (e.g., Asteroxylon, Sphenophyllum). {C] Plectostele A protostele in which the central xylem core breaks into more or less parallel plates is known as plectostele, Each xylem plate is surrounded by phloem (Fig. 1C). Aerial shoots and cone axis of Lycopodium clavatum and L. volubile have this type of stele. {D] Mixed Protostele Sometimes the solid xylem core of the protostele is broken into small groups of tracheids which remain embedded in the phloem (Fig. 1D). Such 4 protostele is known as mixed protostele and is found in the stems of Lycopodium cernuum. A mixed protostele is different from a protostele with mixed pith, The latter possesses groups of Wrachcids intermixed with parenchyma cells (€g, Osmunda regalis, Fig. 2). Siphonostele A ‘simple protostele is sufficient to meet the Tequirements of a small stem or a few larger ones under certain circumstances. However, the Tequitement of increasing diameter of the stem is met with by certain internal modifications in the stele. The first step in this direction is the appearance of definite parenchymatous pith or medulla in the centre of the protostele. Such a modified protostele with a central pith is known as siphonostele or medullated stele, It is an advancement over protostele. Jeffery (1910), on the basis of their association with leaf and branch traces, recognised two types of siphonosteles — cladosiphonic and phyllosiphonic, A cladosiphonic stele _ is characterised by the absence of leaf traces. Microphyllous lycopsids are the main example of this group. Phyllosiphonic stele, on the other hand, has both leaf and branch traces and is found in the members of the order Filicales. Siphonosteles of Equisetum, Osmunda, Schizaea, etc., have a single phloem ring external to the xylem. Such a siphonostele is called ectophloic siphonostele (Fig. 3 A). Contrary to this, the siphonosteles of Adiantum, Dryopteris and Marsilea have a ring of phloem each external and internal to the xylem. This type of siphonostele is called amphiphloic siphonostele (Fig. 3 B). Such steles characteristically have two endodermal layers: outer endodermis — that lies outside the outer phloem and inner endodermis — that lies inner to the inner phloem, outside the pith, [A] Origin of Siphonostele ‘As mentioned earlier, siphonostele has originated by the development of pith in the centre of the protostele. The process of medullation of protostele has been debated, The following two views have been put forward regarding the origin of pith in protostele. [1] Intra-stelar origin of pith ‘According to this view, pith originated as a result of transformation of tracheary elements of the central xylem core into parenchyma. Thus the pith is wholly intra-stelar. Examples of Borrychium ternatum, B. virginianum, Osmunda _regalis, Lunularia, etc, are cited in support of this view where tracheids are scattered throughout the pith (PTERIDOPHYTA)Stelar System i 240 ys In Pleridophyy, ee eee outer endodermis. : ‘outer perioyclo. S outer phloem pith xylem: Inner phloem Inner pericycle inner endodermis A Fig. 3 A+B. Types of siphonosteles : A. Ectophloic siphonostele, B. Amphiphloic siphonostele. cortex Suter endoderm. ‘outer pericycle. leaf gap ‘outer phloem xylem. leaf trace FD teat gap innenchioem Toaf trace Inner paricycle inner endodermis: A B c Fig. 4 A-B. Types of solenosteles : A. Ectopbloic solenostele, A. Amphiphloic solenostele. (ie, mixed pith). This view was supported by [B] Modifications of Siphonostele Boodle (1901), Gwynne-Vaughani (1908), Bower (1911) and Fahn (1960). (IJ) Extra-stelar origin of pith According to this view, protostele has transformed into a siphonostele due to the migration of cortical cells into the stelar axis (Jeffery, 1902, 1910, 1917), Openings, such as leaf and branch gaps, probably provided passage for the invasion of parenchyma, This view derives support from amphiphloic siphonostele that has two endodermal layers, one delimits the stele from the cortex and the other from the pith. Since endodermis is a structure peculiar to the cortex, pith is considered as cortical in origin. (PTERIDOPHYTA) Depending on the presence of non-overlapping ot overlapping gaps, the following three types of siphonosteles are recognised. [1 Solenostele A siphonostele with non-overlapping leaf gaps is known as solenostele, The solenostele may be ectophloic or amphiphloic (Figs. 4A-B, 5A). Usually in the lower part of the stem where leaves are sparsely placed, the stele is solenostelic. [1] Dictyostele Many ferns like Dryopteris, Pteris, Ophioglossum tc,, have a very small thizome with crowded leaves, Consequently, the leaf gaps overlap withaf traces B Fig. 5 A-B. Three-dimensional views of the vascular + A. A solenostele, B. A dictyostele (note overlapping leaf gaps). eo re Fig 6 A-B. Dictyostele : A. As seen in transverse section (cote meriseles), B. Vascular skeleton of a dictyostee, each other (Fig. 5 B). A siphonostcle with overlapping leaf gaps is known as dictyostele, meristeles of ‘outer ring, meristeles of inner ring thick walled ‘ground tissue Fig. 7, Polycyclic stele, (A dictyostele has many scattered vascular strands, each of these strands is called a meristcle (Fig. 6 A, B). {III} Polycyclic stele Stelar structure of certain pteridophytes is still more complex. They possess two or more concentric rings of vascular tissue and are called polyeyclic (Fig. 7). For instance, there are two rings of vascular tissue in Preridium aquilinum, three in Montania pectinata and four in Pteris podophylla and Platycerium aethiopicum. If in a polycyclic stele the outer cylinder is solenostelic, it is called polycyclic solenostele (Fig, 8 A) and if the ower cylinder is dictyostelic, it is known as polycyel (Fig. 8 B). ¢ dictyostele (PTERIDOPHYTA)242 ¥ telar System iy Pleridophytey Important i 7 epOrin Questions 1. Describe the different types of steles present in various members of pteridophytes studied by you With the help of labelled diagrams describe different types of stelar system found in pteridophytes What is stele? Write the evolution of stelar system in pleridophytes, What is a stele? Describe the stclar system of pteridophytes, Waite short notes on {) Siphonostele(i) Plectostele, (i) Actinostele, (iv) van Tieghem and Douliot, (v) Protosele, (vi) Dityostele What is stele? Describe various types of steles found in ptcridophytes with the help of diagrams. Waite an essay on stelar system of pteridophytes. With the help of suitable diagrams, describe the various stelar systems in pteridophytes studied by you What is stele? Give an illustrated description of the various types of steles found in pteridophytes ‘What are cladosiphonic and phylosiphoic siphonostles? Write an account of the origin of siphonostle in peidphje >> Long an Senne Y v Short answer questions Differentiate between protostele and siphonostele. Write a note on origin of siphonostele, Explain how the pith originated in the stele of pteridophytes, How will you differentiate a solenostele from that of a dictyostele? Differentiate a polycyctic solenostele from that of a polycyclic dictyostele. Give a brief account of the various types of protostele found in pleridophytes, Write the main features of stelar theory as proposed by van Tieghem and Douliot (1886). “The term vascular cylinder is better suited than stele in relation to higher plants’. Comment upon the statemen. Name a medullated stele found in pteridophytes. Write a brief account of origin of such a type of stele How will you differentiate between an ectophloic siphonostele and an amphiphloic siphonostele? Write an example for ‘each in which these stcles are found, >> Very short answer questions Name one pleridophytic plant in which amphiphloic siphonostele is found. Name the type of stele in pteridophytes where vascular cylinder is medullated. Which type of stele is found in Marsilea rhizome? What is solenostele? Give one example of dictyostele In which plant protostele is found? In which pteridophyte sctinostele is found? Rearrange the following types of steles in the order of their evolution: siphonostele, plectostele, haplosele, acinstl dictyostele. 9. There are a number of component bits in the dictyostele. What do you call each one of them? 10, What do you call a solenostele which has overlapping leat gaps? 11. What do you call a siphonostele which has developed a few leaf gaps? 42, What is the name given to the protostele which is broken up into mesh like masses, and when the phloem is found fin the interstices? 13. When a medulla of parenchyra is developed in the centre of the protostele, what is it known 14, What is the type of stele found in the simplest vascular crypiogams ? 15. What do you call a stele when the xylem is ridged, exarch and the phloem occupies the bay areas? 16, The vasculature of a higher crypiogam has a core of soft xylem surrounded by phloem, and without leaf gaps "hit is it called? 17. Give an example where you find the steles arranged in many rings 18. Who proposed the stelar theory in pleridophytes? 19. What is the technical term for a non-medullated stele? 20, Name species of Lycopodium where plectosile is found. Py ewaupe >> Fill in the blanks 1. Plectoxtele is prevent in 2, Amphiphilolc siphonostele ix found i i 3. A siphonostele with leaf and branch gaps is culled (PTERIDOPHYTA)Pieridophytes gtr system in . sextet a cena Sli MASS of xylem sue Aula supply NEN Out foe Tea fom them 5 Tm Migg to van Teghen and Dott, the iO 6 ia thse of Marlen the vascular einer ian, OF Yasar ongaiation tw Mle in which the central xylem core breaks ina $A seonastele with non-overlapping leaf gaps is known Piet score OE BiMitag CRT emer Ww PrMpasnhonic stele is characterized by the... f xylem, surrounded by a band of phloem. oof leaf traces, {A aiaystle has many seatered vascular stands, etch of these stands fe known as “d by phloem and erodermis is called ain stele is called ‘ore oF Less parallel plates is known as as false statements Sp Troe and af stele is a non-medullated stele ‘A sphonostele with phloem extemal as well as intemal to xp in lower vascular cryplogams like Lycopodium and Selaginella, independently. the stele being mainly cauline. The prtotele is considered to be the most primitive, both phylogenetically as well as ontogenctcaly, 1n Flcales, provoscle is seen generally in the sporelings. {A proostele in which the central xylem core breaks into mote or less parallel plates is known as actinostele A mixed protostele is different from a protostele with mixed pith; the latter possesses groups of tracheids intermixed swith parenchyma cells. Phpllosiphonic stele has both leaf and branch traces. ‘According to extra-stclar origin, the pith originated as a result of transformation of tacheary elements of the central xylem core into parenchyma. 0. A siphonostele with overlapping leaf gaps is known as a solenostele, is extophoie. the stele of the axis and the leaf traces develop wee > Multiple choice questions 1. Amphiphloic siphonostele is found in the rhizome of: 6. Which of the following does not have a pith? (a) Equseron (0) Marsilea (@) protostele (6) solenostele (6) Selaginella (@) none of the above (©) siphonostele (@)dictyostele 2 Actinostele is found in: 7. Mixed protostele is found in: (2) Lycopodium cermuum(o) Lycopodium clavatum (@) Selaginella (©) Lycopodium (©) Lycopodium serrarum(@) Lycopodium phlegmaria (©) Equisetum (@) Marsitea 3. A stele without a central pith is called: 8. Who proposed the stelar theory? (2) protosele (©) siphonostele (e) Druery (©) Whitier and Steeves (6) solenostele (@) dictyostele (© Zimmermann 4, Actinosile is related to: (@) Yan Tieghem and Doutiot (2) dictyostele (©) siphonostele 9. Dietyostele is found in: (6) solenoscle (© protostele @ Dryopteris (©) Preris 5. Siponosile is found in the stem of: (©) Ophioglossum (dal of the above (6) Pinus (©) Ephedra 10, The vascular strands of a dictyostele are called: (6) Marsilea (@) meristeles (©) telomes (@) all of them (©) apospory (@) apogamy ANSWERS »>Very short answer questions 4. Manilea, 2, Sipbonostele, 3. Amphiphloic siphonostele, 4. A siphonostle with non-overlapping leaf gaps, 5. Previs, 6. Scloginela, peers airy deplete, aciosie,pecosele,sghonostel,dictyosee, 9. Merivee, 10 Diciyostele, 11, Solenostele, 12. Mixed prtostele, 13. Siphonosele, 14, Potoxel, 18. Actinostle, 16. Haplostele, 17. Preridiw aquilinum, 18, van Tieghem and Douliot (186), 19. Protostele, 20. Lycopodium ¢lavanun, >> Fill in the blanks 1 Geapoian clavann, 2. Mae, 3. enone 4. pts 5. kal inet 6 se 7, amphi siphon z 9 solenostele, 10. non-medullated, 11. absence, 12 meristele, >> True and false statements 1. Fae, 2 Fale, 3, True, 4, True, 5. Tru, 6. False, 7, Tre, 8 Trve, 9: Flac, 10. Fase >> Muliple choice questions 1A, 246, 3, 4), 5 (6), 6, (0, 7.) 8 (Ds 9 Dh 10.0)