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VARIATION OF MICROMORPHOLOGICAL CHARACTERS OF LEMMA AND PALEA IN AEGILOPS KOTSCHYI AND AEGILOPS BIUNCIALIS × SECALE CEREALE HYBRIDS, AMPHIPLOIDS AND PARENTAL FORMS

MAŁGORZATA KLIMKO, HANNA PUDELSKA, BARBARA WOJCIECHOWSKA, WOJCIECH KLIMKO

M. Klimko, W. Klimko, Department of Botany, Poznań University of Life Science, Wojska Polskiego C, - Poznań, Poland, e-mail: [email protected] H. Pudelska, B. Wojciechowska, Institute of Plants Genetics, Polish Academy of Sciences, Strzeszyńska /, - Poznań, Poland

(Received: February , . Accepted: March , )

ABSTRACT. Results of this study provided data on new qualitative and quantitative characters of the lemma and palea epidermis in amphiploids of Aegilops kotschyi, Ae. biuncialis × Secale cereale and their parental species using scanning electron microscopy. Several micromorphological characters of both long and short cells (cork cells, crown cells, prickles and macrohairs), stomata and their types, wax and the cuticle were observed. All characters were treated as separate, although crown cells, prickles and macrohairs are treated as single groups (exodermic cells) owing to their protrusion from the periclinal wall. All observed micromorphological characters of lemmas and paleae in amphiploids are inherited after Ae. kotschyi and Ae. biuncialis, irrespective of the method in which these hybrids were produced. Hexaploid (n = x = , x = ) amphiploids in many features exhibit an extended range of variation in comparison to their parental species. In amphiploids in the palea a new shape type is observed in crown cells wide and oblate, not found in either parent form. Moreover, certain dependencies were observed between the method in which am- phiploids were produced and their micromorphological characters. The genera Aegilops and Secale cereale and their amphiploids may be identifi ed using the micromorphological characters of lemmas and/or paleae.

KEY WORDS: Aegilops kotschyi, Aegilops biuncialis, amphiploids, lemma, palea, micromorphology, Secale cereale, SEM

INTRODUCTION and paleae has not been widely studied in the Poideae, except for some particular genera, such as Stipa (THO- Intergeneric hybrids are interesting in the theoreti- MASSON ), Melica and selected species of Briza, cal aspect in evolutionary, taxonomic and cytological Catabrosa, Glyceria, Neostapfi a, Pleuropogon and Schi- studies. Cytological analysis proved that the U genome zachne (THOMASSON ), Bromus (ACEDO and LLA- of Aegilops sp. polyploids is very closely related to the MAS ), Koeleria (KLIMKO and CZARNA ) and U diploid genome of Aegilops umbellulata and it has Calammophila (KLIMKO et AL. ). not been substantially modifi ed during their evolution, Investigations presented in this paper are a continu- whereas the S genome of Ae. kotschyi (UUSS) and the ation of studies on amphiploids of Ae. kotschyi, Ae. biun- M genome of Ae. biuncialis (UUMM) have undergone cialis × Secale cereale and their parental species. To date substantial changes (KIMBER et AL. , KIMBER and studies have been conducted on the morphology and YEN , YEN and KIMBER ). For these reasons proteins of the pollen coat and protoplast in amphip- combinations of two diff erent genomes of Aegilops sp. loids of Ae. kotschyi × Secale cereale (KALINOWSKI et AL. with the Secale cereale genome (RR) seemed to be an in- ), leaf isoenzymes (KALINOWSKI and WOJCIECHOW- teresting material for molecular and micromorphological SKA ) and peptides of pollen grain in Ae. biuncialis investigations. × S. cereale (KALINOWSKI et AL. ). The spikelet and Micromorphological characters are valuable for its parts are considered as diagnostic characters which systematic studies in the family Poaceae. For example, show a wide variation between taxa, especially in rela- the absence of microhairs is characteristic in Pooideae tion to the lemma and palea. (CLAYTON and RENVOIZE ). Micromorphological The aim of the present report was to conduct de- characters of fl oral bracts in grasses have been used to tailed observations of micromorphological characters assess systematic relationships, as well as evolutionary of lemmas and paleae in Ae. kotschyi and Ae. biuncialis trends. Nevertheless, the micromorphology of lemmas × S. cereale amphiploids and their parents. M. Klimko ...

MATERIAL AND METHODS LM. Samples were sonicated in xylene for at least thirty minutes to remove epicuticular wax that may obscure Plant material consisted of the parental species form surface features (ACEDO and LLAMAS ). In this of Ae. kotschyi (accession , in this paper referred paper we followed ELLIS () for the description of to as AK-), Ae. biuncialis (accession , referred lemma micromorphology, since lemmas are homolo- to as Ab ), and the S. cereale self-compatible line (WS gous to leaves (SNOW ), and extended the termi- N/, denoted as S ). nology to paleae, because they show similar epidermal Amphiploids C (Ae. kotschyi × S. cereale) and A characteristics. (Ae. biuncialis × S. cereale) were produced by colchicine The shortest Euclidean distances were calculated in treatment of F hybrids. Amphiploids E (Ae. kotschyi an agglomerative grouping by the nearest neighbour × S. cereale), and A (Ae. biuncialis × S. cereale) were method and dendrograms were constructed to exam- obtained by chromosome doubling via in vitro culture ine the relationships between amphiploids and parental propagation (WOJCIECHOWSKA and PUDELSKA ) species (SOKAL and ROHLF ). Material was collected from greenhouse-grown plants from the Institute of Plant Genetics, Polish Acad- emy of Sciences in Poznań. The lemmas and paleae were RESULTS analysed in the lowest fl oret of the spikelet in the middle parts of the spike. Data were gathered from the middle Description of micromorphological characters part of the lemma (abaxial surface) (Fig. A), while due Long cells to the considerable variation of characters in the paleae Long cells are the dominant element in the epider- near the apex, observations were conducted on the mid- mis of lemmas and paleae, which, despite their name, dle part and along veins (Fig. B). vary notably in length. There are rectangular, long and Observations were carried out fi rst with a stereo mi- narrow cells having convex periclinal walls. In contrast, croscope, a light microscope (LM) and fi nally a scan- the anticlinal walls are parallel and highly sinuous, with ning electron microscope (SEM). Stains such as Sudan W-shaped waves. The ends are often angular, ranging III were used to aid the identifi cation of cork cells using in length from . to . μm. Short cells Short cells are in general nearly equal-sided, al- though cells with the width exceeding their length are also relatively frequent. Short cells alternate with long cells, or occur solitarily or in pairs. The frequency of short cells results to some degree from the length of long cells and shows variations between species and amphiploids. Anticlinal cell walls are straight or sinu- ous. Two types of short cells have been observed: cork cells and exodermic cells. Cork cells They are short cells containing solid deposits of or- ganic substances and suberifi ed cell walls (KAUFMAN et AL. ). Their periclinal walls are usually slightly con- vex and collapse upon dehydration. The most frequent shape is elliptic, reniform and semicircular. Exodermic cells These cells protrude beyond the general surface of the epidermis and have periclinal walls with diff ering degrees of convexity. Depending upon the size and shape of the processes, three subtypes may be distinguished: Crown cells: short exodermic cells with small con- vex or conical protrusions, which occur frequently on the lemma and palea. This term was used by PRAT () and revived by WATSON and DALLWITZ () in order to characterize the leaf epidermis in some genera of grasses. Many authors include all exoder- mic cells under the name of macrohairs. ELLIS () referred to these cells as hooks, using the term as a synonym of crochets and crown cells, although he only presented fi gures with hooked protrusions, thereby excluding those with rounded protrusions. The protrusions of the periclinal wall of crown cells vary in shape and size. In general they have crenu- late walls, with pronounced waves similar to those FIG. . Observation area: A – lemma, B – palea (abaxial of long cells. surface) Variation of micromorphological characters of lemma and palea in Aegilops kotschyi ...

· Prickles: Exodermic cells with elongated tips and cells per mm. Crown cells rounded with a spherical swollen bases arising directly from the epidermis. and conical protrusion (Fig E), density - crown They occur frequently especially on veins and the cells per mm. Prickles, macrohairs in the middle part marginal zone, although in some groups they occur absent. Macrohairs present very rarely at the margins between long cells with a comparable frequency, and and at the base of the lemma, .-. μm long. Sto- a high silica concentration in this structure. mata .-. μm long. Wax dense, fi liform, covering Macrohairs: Unicellular structures, soft or rigid, almost all the surface, the cuticle irregularly, longitu- long, often with a bulbous base. They occur between dinally striated and nodular (along anticlinal walls of other epidermal cells, the marginal zone and in some long cells (Fig. F). cases they have an associated small cork cell. Palea. Long cells .-. μm. Cork / silica cell Epicuticular wax – pairs reniform / elliptic .-. μm long, density Epicuticular waxes are deposits, formed mainly by - cork cells per mm. Crown cells rounded, .- a large mixture of diff erent chemical compounds. -. μm long, density - crown cells per mm. Crown cells in the middle part of the surface diff er Micromorphological description of lemmas and paleae considerably in terms of their shape: with conical and pointed protrusions (Fig. F). Prickles and macrohairs Aegilops kotschyi (AK-) distributed mainly in the marginal zone. Stomata .- Lemma. Long cells .-. μm. Cork / silica – cell -. μm long, wax absent, the cuticle smooth, striated pairs reniform / elliptic .-. μm, density - cork transversely and longitudinally striated, nodular (along cells per mm. Crown cells rounded, .-. μm anticlinal walls of long cells and stomata) (Fig. G). long, with a conical protrusion (Fig. A), density - crown cells per mm. Prickles in the middle part, Aegilops kotschyi × S. cereale ( C) near the apex and the marginal zone absent or scarcely Lemma. Long cells .-. μm. Cork / silica – distributed mainly on veins. Macrohairs - mm-, up to cell pairs reniform / elliptic, .-. μm, density - . μm long. Stomata .-. μm long. Wax granu- - cork cells per mm. Crown cells rounded .-. late, the cuticle smooth and irregularly, longitudinally μm long, with a conical protrusion (Fig. A), density striated (Fig. B). - crown cells per mm. Prickles in the mid- Palea. Long cells .-. μm. Cork / silica cell dle part absent; single, wide prickles present on veins. – pair reniform / elliptic .-. μm, density - Macrohairs absent. Stomata .-. μm long. Wax cork cells per mm. Crown cells rounded .-. μm, dense, granulate, covering almost all the surface. The density - crown cells per mm. Crown cells in cuticle striated longitudinally, masked by wax (Fig. B). the middle part rounded (Fig. A), with a conical pro- Palea. Long cells .-. μm. Cork / silica cell – trusion. Closer to the apex a considerable prevalence of pairs reniform / elliptic .-. μm, density - crown cells with pointed protrusions (Fig. B). Prick- cork cells per mm. Crown cells rounded .-. les present at the margin and in the area of the apex. μm long with conical and pointed protrusions (wide and Macrohairs absent in the abaxial epidermis. Stomata oblate) (Fig. A), density - crown cells per mm. .-. μm long, wax absent, the cuticle smooth and This is a new character. Prickles and macrohairs absent nodular (along anticlinal walls of long cells) (Fig. C). in the middle part, found only on veins and crenations and the margin of the palea. Stomata .-. μm Secale cereale (S ) long. Wax granulate, irregularly distributed, scarce. The Lemma. Long cells .-. μm. Cork / silica – cuticle fi nely, longitudinally striated and nodular along cells pairs semicircular / elliptic .-. μm, density anticlinal walls of long cells (Fig. B). - cork cells per mm. Crown cells rounded, .-. μm long, with pointed protrusions (Fig. C), Aegilops kotschyi × S. cereale ( E) density - crown cells per mm. Prickles generally Lemma. Long cells .-. μm. Cork / silica absent in most of the lemma, sometimes several prick- – cell pairs reniform / elliptic .-. μm, density les may be found on veins. Macrohairs absent. Stomata - cork cells per mm. Crown cells rounded and .-. μm long. Abundant accumulations of mixed irregular .-. μm long, with conical protru- wax (fl aky-fi liform), masking the cuticle, which is stri- sions, density - crown cells per mm. Not very ated transversely and longitudinally (Fig. D). numerous prickles on middle veins (Fig. C) and the Palea. Long cells .-. μm. Cork / silica cells marginal zone. Macrohairs absent. Stomata .-. – pairs semicircular / elliptic .-. μm long, den- μm long. Wax granulate, abundant. The cuticle smooth, sity - cork cells per mm. Crown cells rounded transversely striated and nodular along anticlinal walls with a pointed protrusion (Fig. D), generally absent of long cells and stomata (Fig. D). in most of the palea, although sometimes several may Palea. Long cells .-. μm. Cork / silica – cell be fund near the base or beside the veins. Macrohairs pairs reniform / elliptic .-. μm long, density absent. Stomata .-. μm long. Wax mixed (granu- - cork cells per mm. Crown cells rounded and late-fi liform), abundant. The cuticle smooth and striated irregular .-. μm long, with conical and pointed longitudinally (Fig. E). protrusions, cauched (Fig. C), density - crown cells per mm. Prickles in the middle part absent, Aegilops biuncialis (Ab ) distributed mainly do / length of the palea on veins. Lemma. Long cells .-. μm. Cork / silica – cell Stomata .-. μm long. Wax granulate, the cuti- pairs reniform / elliptic .-. μm, density - cork cle smooth, irregularly, longitudinally and transversely M. Klimko ...

FIG. . SEM. Lemma surface: A, B – Aegilops kotschyi (AK-); C, D – Secale cereale (S ); E, F – Aegilops biuncialis (Ab ) Variation of micromorphological characters of lemma and palea in Aegilops kotschyi ...

B

FIG. . SEM. Palea surface: A, B, C – Aegilops kotschyi (AK-). Note A in the middle part and B near the apex; D, E – Secale cereale (S ), F, G – Aegilops biuncialis (Ab ) M. Klimko ...

FIG. . SEM. Lemma surface of amphiploids: A, B – Aegilops kotschyi × Secale cereale ( C); C, D – Ae. kotschyi × S. cereale ( E); E, F – Ae. biuncialis × S. cereale ( A); G, H – Ae. biuncialis × S. cereale () Variation of micromorphological characters of lemma and palea in Aegilops kotschyi ...

FIG. . SEM. Palea surface of amphiploids: A, B – Aegilops kotschyi × Secale cereale ( C); C, D – Ae. kotschyi × S. cereale ( E); E, F – Ae. biuncialis × S. cereale ( A); G, H – Ae. biuncialis × S. cereale () M. Klimko ... striated and papillated along anticlinal walls of long Aegilops sp. For this reason observations were conducted cells (Fig. D). near the apex and along veins. On the basis of the epiderma micromorphology of Aegilops biuncialis × S. cereale ( A) the lemma and palea in the studied taxa and Ae. variabi- Lemma. Long cells -. μm. Cork / silica – lis, Ae. ovata, Ae. cylindrica (KLIMKO, unpublished data) cells pairs or solitary reniform / elliptic, .-. μm it may be inferred that species of the genus Aegilops long, density - cork cells per mm. Crown cells have the same epidermal cell types, diff ering markedly rounded or irregular .-. μm long, with conical from those in S. cereale. In Aegilops and Secale there are and pointed protrusions (wide, oblate), cauched (Fig. two characters that in general show a positive correla- E) density - crown cells per mm. Prickles distrib- tion. They are the length of long cells (in the lemma and uted mainly on veins and awns. Stomata .-. μm palea) and the number of short cells. More diff erences long. Wax mixed (granulate–fl aky), irregularly distribut- occur in short cells than long ones. Thus, crown cells ed. The cuticle smooth and fi nely striated, nodular along show considerable variation in density and their type of anticlinal walls of long walls and stomata (Fig. F). protrusions, whereas cork cells exhibit variation in their Palea. Long cells .-. μm. Cork / silica cells density and size. Exodermic cells, such as e.g. cork cells, – pairs, reniform / elliptic .-. μm long, den- are distributed in longitudinal patterns, alternating with sity - cork cells per mm. Crown cells rounded or long cells and other epidermal cells. The density of cork irregular .-. μm long, density - crown and crown cells in relation to the other epidermal ele- cells per mm .Crown cells with conical and pointed ments is also variable. protrusions (wide, fl attened), cauched (Fig. E). Prickles Lemmas of parental species Ae. biuncialis and Ae. and macrohairs distributed mainly in the marginal zone. kotschyi diff er in terms of the density of cork cells per Stomata .-. μm long. Wax granulate, accumu- mm, which amounts to . and ., respectively. lated mainly around cork cells. The cuticle smooth, lon- Much bigger interspecifi c diff erences were observed in gitudinally striated and nodular along anticlinal walls the density of crown cells per mm, which was in of long cells and stomata (Fig. F). Ae. kotschyi and . in Ae. biuncialis. No signifi cant diff erences were observed in this trait in the palea. Aegilops biuncialis × S. cereale () Secale cereale (S) is characterised by a consider- Lemma. Long cells .-. μm. Cork / silica able prevalence of the density of cork cells per mm, – cell pairs reniform / elliptic – oval, .-. μm amounting in the lemma to and in the palea to long, density - cork cells per mm. Crown cells . cells, respectively. The density of crown cells per rounded .-. μm long, with conical protrusions mm in the lemma is ., while in the palea it is only (Fig. G), density - crown cells per mm. Prickles .. distributed mainly in the marginal zone and on awns. Crown cells are generally rounded, although some- Stomata .-. μm long. Wax mixed (granular– times they are irregular in shape as in the palea of fl aky-fi liform), abundant. The cuticle smooth, longitu- amphiploids and A. The protrusion or protuber- dinally striated and nodular along stomata (Fig. H). ance may be conical or pointed and vary in its relative Palea. Long cells .-. μm. Cork / silica cell size to the cell supporting it. The protrusion is formed – pairs, reniform / elliptic .-. μm long, density by most of the external periclinal wall of the cell. - cork cells per mm. Crown cells rounded or ir- The density of cork cells per mm in Ae. kotschyi, regular .-. μm long, with spherical and pointed Ae. biuncialis and S. cereale is higher in the palea than protrusions (wide, fl attened), cauched (Fig. G), density the lemma. In amphiploids the trend is opposite. In - crown cells per mm. Prickles and macrohairs AK- and S the density of crown cells per mm is present on veins and in the marginal zone. Stomata higher in the lemma than the palea. Ab and all am- .-. μm long. Wax granulate, scarce. The cuticle phiploids have highest values for the mean number of transversely and longitudinally striated, nodular along crown cells per mm in the palea. Amphiploid C in anticlinal walls of long walls (Fig. H). comparison E has a higher density of cork and crown cells per mm in the lemma and palea. In amphiploid A in comparison to only the density of crown DISCUSSION AND CONCLUSIONS cells in the lemma and palea is higher. Prickles on the lemma in studied taxa occur especially above the veins. One of the main problems with micromorphologi- Long macrohairs in amphiploids and parent species were cal studies is to determine the optimal site of observa- found very rarely in the abaxial epidermis. Their pres- tion. The lemma shows variations in the expression of ence or absence is of little taxonomic signifi cance. It is micromorphological characters between the apex, base in contrast to the glume, where the type of prickles in and margins, whereas the palea varies mostly near the Ae. kotschyi (semi-erect prickles), Ae. biuncialis (cauched apex and the marginal zone. Thus the middle part of prickles) and their distribution eff ectively diff erentiate the lemma side and the central part of the palea body both species (MHAIDAT et AL. ). were selected for analyses. These regions appear to be All studied taxa have macrohairs and prickles re- quite constant in their micromorphological charac- stricted to the margin of the palea. Sometimes prickles teristics when studied in diff erent taxa. However, this are longer and then it is diffi cult to diff erentiate between area does not fully illustrate the entire variation in the prickles and macrohairs (METCALFE , ELLIS , analysed micromorphological characters in the palea of SNOW ). Variation of micromorphological characters of lemma and palea in Aegilops kotschyi ...

The size of stomata is considered to be an indirect biuncialis) and their amphiploids. In terms of analysed indicator in studies on polyploidy (WINKELMANN and characters S. cereale showed the biggest diff erences GRUNWALDT ). On the lemma and palea stomata (S ). are arranged in rows along veins. The mean length of On this basis the following conclusions may be in- stomata in the abaxial epidermis of the lemma ranged ferred. For the micromorphological studies it is neces- from . μm (Ae. biuncialis) to . μm (Ae. kot- sary to carefully select the site for observations. All schyi) and . μm (S. cereale). The length of stomata micromorphological characters in the lemma and palea in amphiploid C was . μm. Stomata in amphip- in the amphiploids are inherited after Ae. kotschyi or Ae. loid E were much shorter, of . μm. The length biuncialis, irrespective of the method in which they were of stomata in amphiploids A and was . μm produced. Hexaploid amphiploids in terms of analysed and . μm, respectively. characters exhibit extensive variation in comparison to In the palea mean diff erences in the analysed trait the parental species. Moreover, near the apex on the between parental species were smaller, as the value palea of amphiploids a new shape of crown cells was was . μm (Ae. kotschyi), . μm (Ae. biuncialis) observed, a wide and oblate, not found in the parental and . μm (S. cereale). The length of stomata in am- species. A certain variation was also found in the rela- phiploid C was . μm, while it was . μm in tionship between the method of amphiploid production amphiploid E. The mean value of this character in (colchicine treatment or in vitro propagation) and their amphiploids A and (Ae. biuncialis × S. cereale) micromorphological characters, similarly as it was the was . μm and . μm, respectively. case in studies on pollen grain sculpture (KALINOWSKI In studied taxa the length of stomata was higher in et AL. ). Selected species of Aegilops, Secale cere- the lemma than the palea in the parental species and ale and their amphiploids may be identifi ed using the their amphiploids, except for amphiploid E, where micromorphological characteristics of the lemma and/ these diff erences were slight. The interesting aspect or palea. of the epicuticular wax residues, according to some authors, lies in the fact that the confi guration of wax deposits seems to have a genetic basis (BAUM and HA- REFERENCES DLAND ). On the lemma the morphology is diff erent in Ae. kotschyi, S. cereale and Ae. biuncialis (Figs B, ACEDO C., LLAMAS F. (): Variation of micromorpho- D, F). In amphiploids C, E and A wax granu- logical characters of lemma and palea in the genus late is found, in mixed (granular-fl aky-fi liform). In Bromus (Poaceae). Ann. Bot. Fenn. : -. analysed taxa granular wax predominated on the palea, BARTHLOTT W., NEINHUIS C., CUTLER D., DITSCH F., in S it was mixed, while in AK- and Ab it was not MEUSEL I., THEISEN I., WILHELMI H. (): Classi- observed. Detailed observations of epicuticular wax on fi cation and terminology of plant epicuticular waxes. the abaxial surface provided new data on the subject. Bot. J. Linn. Soc. : -. The cuticle on the lemma and palea may be smooth, BAUM B.R., HADLAND V.E. (): Epicuticular waxes papillated and striated (nomenclature following BAR- of glumes of Avena: a scanning electron microscopic THLOTT et AL. ). The cuticule sculpture in the study of the morphological patterns in all the spe- lemma and palea of the parental species and amphip- cies. Can. J. Bot. : -. loids did not vary. In contrast, marked diff erences were observed in a thick cuticle layer of the culm: smooth in CLAYTON W.D., RENVOIZE S.A. (): Genera Grami- Ae. kotschyi and striated in Ae. biuncialis (MHAIDAT et num: Grasses of the world. Royal Bot. Gard. Kew Bull. AL. ). Addl. Ser. . Cluster analysis using Euclidean distances segregat- ELLIS R.P. (): A procedure for standardizing com- ed all the taxa compared within the one group (Fig. ). parative leaf anatomy in the Poaceae II. The epider- It was formed by parental species (Ae. kotschyi, Ae. mis as seen in surface view. Bothalia : -. KALINOWSKI A., KLIMKO M., WOJCIECHOWSKA B. (): Morfologia oraz białka płaszcza i protoplastu pył- ku u amfi ploidów Aegilops kotshyi × Secale cereale. AK-3 In: Genetyka w ulepszaniu roślin użytkowych. Eds 84 C P. Krajewski, Z. Zwierzykowski, P. Kachlicki. Instytut Genetyki Roślin PAN, Poznań: -. 84 E KALINOWSKI A., WOJCIECHOWSKA B. (): Two-di-

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S 14 cialis × Secale cereale hybrids and Ae. kotschyi × S. ce- reale amphiploids in relation to their parents. J. Appl. Genet. , : -. 0 50 100 150 200 250 KAUFMAN P.B., PETERING L.B., SMITH J.G. (): Ul- FIG. . Dendrogram constructed on basis of quantitative trastructural development of cork-silica cells pairs in and qualitative characters of lemma and palea Avena internodal epidermis. Bot. Gaz. : -. M. Klimko ...

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