ࡱ>  HJLN~#`fFbjbj\.\.`R>D>Dg%$pppp( ( ( $L t_t_t_P_`,L )bLc"ncncncnn#o$Go#h ( vx+mnvxvxppncnc 6vxhp<nc( ncvx |( pncb @Pt_ޒ@ H)Xyy0py( p([o`qms\t[o[o[o^[o[o[o)vxvxvxvxL L L $HpR L L L pRL L L pppppp Stomata in some species of genus Arum from the Eastern Slavonia and Baranya region Mirjana Sabo 1, Tihana Mar ek1, Tomislav Ba i2, Ljiljana Krstin2 & Anita Lendel2 1Faculty of Food Technology, J.J. Strossmayer University, 31 000 Osijek, Croatia 2Faculty of Philosophy, Department of Biology, J.J. Strossmayer University, 31 000 Osijek, Croatia Types and the number of stomata in the following Arum species: Arum italicum Mill., Arum maculatum var. maculatum L. and Arum maculatum var. immaculatum L., Arum alpinum var. pannonicum Terpo. and Arum alpinum var. intermedium Schur. at three different locations in Zablae, Normanci and Bilje were investigated. The most prevalent stomata type at both upper and lower epidermis for each Arum species was mostly stomata type paracytic, followed by hexacytic, tetracytic or brachyparacytic as far as locality is concerned. Helicocytic type was more prevalent in Arum alpinum var. pannonicum Terpo. and Arum alpinum var. intermedium Schur. at Bilje. Other stomata types were very rare. A striking regularity in the occurrence of stomata types was not found within a single species. Some stomata types, however, were found either at adaxial or abaxial epidermis or were not present at all. The number of stomata per square mm varied from 25 to 651. A statistically significant differences in the number of stomata per square mm at upper and lower epidermis among Arum species was determined at locations Zablae and Normanci, whereas no statistically significant differences were found in location Bilje. Keywords: Arum species, stomata types, stomata number ____________________________________________________________________________ Corresponding author: Mirjana Sabo, Faculty of Food Technology, J.J. Strossmayer University, F. Kuha a 18, 31 000 Osijek, Croatia, fax: +385 31 20 71 15, e-mail:  HYPERLINK "mailto:mirjana.sabo@ptfos.hr" mirjana.sabo@ptfos.hr Introduction Metcalfe & Chalk (1979) classify stomata according to their number and the shape of subsidiary cells (anisocytic, anomocytic, brachyparacytic, helicocytic, hemiparacytic, hexacytic, paracytic, staurocytic, tetracytic, and hemidiacytic). Helicocytic stomata type is surrounded by four or more subsidiary cells, which have, according to Payn (1970) spiral thickenings and can be found in Brassicaceae, Crasulaceae and Begoniaceae families. Van Cotthem (1970) described hexacytic stomata type as the one whose guard cells are surrounded with four subsidiary cells. According to Van Cotthem (1970), paracytic type is surrounded by two guard cells. A stoma can cell It is not possible to say that a cell can consist of more cells can consist of one or more vertical elongated subsidiary cells parallel to stoma with longer axis. Staurocytic stomata type was introduced by Van Cotthem (1970) who defined it as 3-5 equal. s equal? subsidiary cells which surround stoma, that have subsidiary cell walls differently oriented with regard to pores. In tetracytic type, guard cells are surrounded with four subsidiary cells; two lateral and two terminal ones. This stomata type is found in many most monocotyledons I am not sure that it is possible to say that tetracytic type is found in most monocots it is not found e.g. in Poaceae, Cyperaceae, Iridaceae and Liliaceae (Metcalfe, 1961). Stomata can also be classified according to their development. Florin (1958) distinguishes perigenous type if there is no common origin of guard and subsidiary cells. The adult stage of this type is also called haplocheilic. The aim of this paper was to determine which was the prevalent stomata type at all locations; secondly, to determine whether Arum species can be differentiated according to the type of stomata and to examine the effect of the habitat on the stomata types. Moreover, the aim was to find statistically significant differences in stomata types among the different species at the same locations and the same species at the different locations too. In addition the aim was to determine whether there were any differences regarding stomata types at upper and lower epidermis and which type was the most prevalent for Arum species. Furthermore, the aim was to determine the number of stomata per square mm at both upper and lower epidermis, and to determine any statistically significant difference in number of stomata among the different species belonging to the same location and among both the same and different species from different habitats. Material and methods The locations chosen differed in area, slightly in height above sea level, phytocenosis, and climatic conditions. Loess soil is typical for each location. The Zablae area consisted of 3. 30 ha, Normanci of 1661.18 ha, and Bilje consisted of 8. 90 ha. The altitude was 89-90 m, 88-93 m, and 91 m. Each location belongs to the same phytocenosis of hornbeam and oak wood Carpino betuli-Quercetum roboris (Raua, 1969). The climatic conditions were mild and pluvial with warm summers and moderately cold winters. Precipitations were equally distributed and winter was the driest period. July was the warmest month with the mean monthly temperature of 21.1 C, in contrast to January, which was the coldest with the mean air temperature of 1.2 C. The mean annual temperature was 10.8 with the mean annual precipitation of 700 mm (data obtained from Croatian Forests, a company for managing forests and forest lands in Croatia). The object of this research was stomata of five species of genus Arum: Arum italicum Mill., Arum maculatum var. maculatum L., Arum maculatum var. immaculatum L., Arum alpinum var. pannonicum Terpo. and Arum alpinum var. intermedium Schur. At Zablae and Normanci locations the following three species were studied: Arum italicum Mill., Arum maculatum var. maculatum L., Arum maculatum var. immaculatum L. At Bilje location the following two species were studied: Arum alpinum var. pannonicum Terpo. Arum alpinum var. intermedium Schur. Determination of stomata type Determination was performed according to Metcalfe & Chalk (1979). Stomata type was determined at ten same leaves of each species. This was the same as for determination of stomata number Does it mean that the same leaves were used for both stomata type determination and stomata number determination? . The leaves of each species at three leaf regions, basal, middle and top leaf part, at upper and lower epidermis, were taken. The upper and lower epidermises were peeled off, cut into short squares, using a fine forceps. Afterwards the samples were put through ethanol series and than stained by alcian blue and safranin. Finally the samples were mounted on microscope slides in glycerin jelly and covered by slips. Determination of the number of stomata per mm2 Stomata were counted randomly at 10 microscopic fields of vision of each mentioned region, from both upper and lower epidermis, in order to obtain the better mean number of stomata per square mm. Therefore, the results according to leaf regions, were not present here. Subsequently the values were multiplied by a beforehand determined surface factor (F=11.6977) and number of stomata were examined at a magnification of 400 x. Surface factor was obtained by standard methods e.g. number of objective lens micrometer marks was multiplied with ten and divided with number of ocular lens micrometer marks. Statistical analyses The results were tested with the t-test in order to determine statistically significant differences for the examined parameters among different Arum species within the same location and also among the same and different species belonging to the two different locations. The t-test was done by means of Microsoft Excel 7 and Origin 6.1, (P0.05, 0.01). Results Stomata types Incidence of stomata types at upper and lower epidermis in Arum italicum Mill., Arum maculatum var. maculatum L., and Arum maculatum var. immaculatum L. at location Zablae was shown in Tab.1. In this location paracytic stomata type (Fig. 1) was the mostly prevalent in each Arum species, at both adaxial and abaxial epidermis (Tab. 1), followed by hexacytic (Fig. 2) and tetracytic (Fig. 3, Tab. 1) or brachyparacytic type (Fig. 4). Brachyparacytic type was mostly on the fourth place at upper epidermis, whereas at lower one was on the third place. Other stomata types, staurocytic, anisocytic, heliocytic, anomocytic, hemiparacytic and hemidiacytic, at this habitat had mostly less than of 0.73 stomata, both at upper and lower epidermis. Incidence of stomata types at upper and lower epidermis in Arum italicum Mill., Arum maculatum var. maculatum, and var. immaculatum L. at location Normanci was shown in Tab. 2. Here paracytic stomata type (Fig. 1) was also mostly prevalent in each Arum species at both upper and lower epidermis, except in Arum italicum L. This type was followed by hexacytic (Fig. 2) and brachyparacytic stomata type (Fig. 4. Tab. 1). According to frequency brachyparacytic stomata type at this location was on the third place (Fig. 3, Tab. 2), whereas tetracytic stomata type was on the fourth place. Other types, from staurocytic to hemidiacytic, were present with less than of 0.60 stomata per leaf, per square mm of leaf.? (Tab. 2, Figs. 5 -10). Incidence of stomata types at upper and lower epidermis in Arum alpinum var. pannonicum Terpo. and Arum alpinum var. intermedium Schur. at location Bilje was shown in Tab. 3. As it was recorded at Zablae and Normanci (Tabs 1 and 2), the most prevalent stomata type in Bilje, at both upper and lower epidermis was paracytic type (Fig. 1), except in Arum alpinum var. pannonicum Terpo.. This type was also followed by hexacytic (Fig. 2) and with brachyparacytic stomata type (Fig. 4). Tetracytic stomata type was here on the fourth place. Other types, from staurocytic to hemidiacytic, were represented with less than 0.35 stomata. Comparison of lower epidermis in Arum alpinum var. pannonicum Terpo. and Arum var. intermedium Schur. revealed that the latter had more paracytic (Fig. 1) and brachyparacytic stomata type (Fig. 4) than Arum alpinum var. pannonicum Terpo. which had more hexacytic (Fig. 2) and tetracytic stomata type (Fig. 3). Concerning upper epidermis of Arum species from each three locations (Zablae, Normanci and Bilje) species had predominantly paracytic type (Fig. 1), followed by hexacytic (Fig. 2) and both tetracytic and brachyparacytic stomata type (Figs. 3, 4), whereas lower epidermis had also predominantly paracytic stomata type (Fig. 1), followed by hexacytic (Fig. 2), brachyparacytic (Fig. 4) or tetracytic types (Fig. 3). Other stomata types, from staurocytic to hemidiacytic, (Figs. 6 to 10) were rather unequally distributed in each Arum species regardless of the location (Tabs. 1, 2 and 3). With regard to this part of results obtained, it can be concluded that differences among locations exist in sequence of stomata types (tertacytic tetracytic type in Zablae and brachyparacytic in Normanci and Bilje). Also, these observations suggest more frequency distribution of stomata type at lower epidermis, but of same stomata type, in contrast to upper ones. This sentence is not understandable, would you formulate it better? Number of stomata per square mm Mean number of stomata per mm2 at upper and lower epidermis in Arum species (t-test) at Zablae and Normanci is shown in Tab. 4. Statistically significant differences between the number of stomata per mm2 at upper epidermis in Arum species at Zablae and Normanci were not found at all (Tab. 4). However, statistically significant differences were determined in the number of stomata per mm2 at lower epidermis between Arum italicum Mill. at location Zablae (235.00 stomata/mm2) and location Normanci (651.00 stomata/mm2) (Tab. 4). With regard to the number of stomata per square mm at lower epidermis statistically significant differences were found between Arum italicum Mill. at locatin Zablae (235.00 stomata/mm2) and Arum maculatum var. immaculatum L. at location Normanci (403.00 stomata/mm2); between Arum maculatum var. maculatum L. at location Zablae (196.00 stomata/mm2) and Arum italicum Mill. at location Normanci (651.00 stomata/mm2); between Arum maculatum var. maculatum L. at location Zablae (196.00 stomata/mm2) and Arum maculatum var. immaculatum L. at location Normanci (403.00 stomata/mm2) and finally between Arum maculatum var. immaculatum L. at location Zablae (290.00 stomata/mm2) and Arum italicum Mill. at location Normanci (651.00 stomata/mm2) (Tab. 4). However, the mean number of stomata per square mm at lower epidermis of all Arum species at locations Zablae and Normanci was greater for all the species at Normanci (453.00 stomata/mm2) than at Zablae (240.3 stomata/mm2) (Tab. 4). The t-test did not reveal any statistically significant differences in the number of stomata per square mm among Arum species at location Zablae (Tab. 4) and Bilje (Tab. 5). However, a statistically significant differences were found in the number of stomata per square mm at location Normanci and Bilje (Tab. 5). Statistically significant difference of p<0.05 was found in the number of stomata per square mm at lower epidermis at location Normanci between Arum italicum Mill. (651.00 stomata/mm2) and Arum maculatum var. maculatum L. (305.00 stomata/mm2) and between Arum italicum Mill. (651.00 stomata/mm2) and Arum maculatum var. imamculatum L. (403.00 stomata/mm2) (Tab.5). The number of stomata per squer square mm at upper epidermis in location Bilje between Arum alpinum var. pannonicum Terpo. was 106.00 stomata/mm2 and in Arum alpinum var. intermedium Schur. 76.30 stomata/mm2. The number of stomata per mm2 at lower epidermis in Arum alpinum var. pannonicum Terpo. and Arum alpinum var. intermedium Schur. totaled 397.00 and 381.00 stomata per mm2 respectively. In other words, Arum alpinum var. pannonicum Terpo had larger number of stomata per squer square mm in location Bile than Arum alpinum var. intermedium Schur. The t-test did not reveal any statistically significant differences in stomata number per mm2 between Arum species at location Bilje (Tab. 5). With regard to this part of the results, e. g. of number of stomata, it can be concluded that number of stomata per square mm varied from 25 to 651. Statistically significant differences in this property at upper and lower epidermis among Arum species were determined at locations Zablae and Normanci, whereas no statistically significant differences were found in location Bilje. Discussion The results of this research indicated that the most prevalent type of stomata in the leaves of the Arum species were paracytic. This is in accordance with Pant et al. (1966) who claimed that paracytic and tetracytic are prevalent stomata types in the family Araceae, followed by hexacytic, tetracytic or brachyparacytic at each location (Tabs. 1, 2 and 3). Hexacytic, as well as tetracytic and brachyparacytic were more prevalent at upper epidermis than at lower epidermis. Helicocytic stomata type is more prevalent in Arum alpinum var. pannonicum Terpo. and Arum alpinum var. intermedium Schur. at Bilje than in Arum italicum Mill., Arum maculatum var. maculatum L. and Arum maculatum var. immaculatum L. at Zablae and Normanci. Other stomata types were rare and unevenly distributed at upper and lower epidermis in each examined species at each location. Guard cells of these Arum species were elongated. In some species guard cells are obtuse, pointed, amputated or rounded. The mean number of stomata at upper epidermis of each species at Zablae amounted 32 stomata/mm2, in contrast to 240.3 stomata/mm2 found at lower epidermis. Furthermore, the mean number of stomata at upper epidermis at Normanci was 31.5 stomata/mm2 and 453 stomata/mm2 at lower epidermis. The values for upper epidermis were approximately equal at both locations, whereas the number of stomata at lower epidermis differed considerably (Tab. 4). Comparison of the same and different species at Zablae and Normanci revealed statistically significant differences in the number of stomata per square mm at upper and lower epidermis between Arum italicum Mill. and Arum maculatum var. immaculatum and var. maculatum L. at lower epidermis and among Arum maculatum var. maculatum and var. immaculatum L. and Arum italicum Mill. at upper epidermis (Tab. 4). At Zablae, in Arum italicum Mill. were recorded at upper epidermis 26.40 stomata/mm2 and in Arum maculatum var. maculatum L. 36.50 stomata/mm2. At the same time, Arum italicum Mill. at Normanci had 24.90 stomata/mm2 and Arum maculatum var. maculatum L. 31.70 stomata/mm2, at upper epidermis. These results indicate that Arum italicum Mill. has larger number of stomata at upper epidermis than Arum maculatum var. maculatum L. at both locations. Furthermore, there were 235.00 stomata/mm2 at lower epidermis in Arum italicum Mill. and 196.00 stomata/mm2 in Arum maculatum var. maculatum L. at Zablae, whereas at Normanci Arum italicum Mill. and Arum maculatum var. maculatum L. had 651.00 stomata/mm2 and 305.00 stomata/mm2 respectively (Tab. 4). In other words, the number of stomata per mm2 at lower epidermis in Arum maculatum var. L. maculatum was lower than in Arum italicum Mill. at both locations. However, at both locations Arum maculatum var. maculatum L. had a greater number of stomata per square mm at upper epidermis than Arum italicum Mill. In comparison to Arum italicum Mill., Arum maculatum var. maculatum L. had a greater number of stomata per square mm at upper epidermis, but number of stomata at lower epidermis at both locations, which proved Saxs theory about less number of stomata at lower epidermis in relation to upper ones (Sax, 1938). Comparison of the species at Zablae and Bilje did not reveal statistically significant differences in the number of stomata per mm2. However, statistically significant differences were recorded at Normanci among lower epidermis in Arum italicum Mill. and Arum maculatum var. maculatum and immaculatum L. and between Arum italicum Mill. A statistically significant difference in the number of stomata per mm2 between Arum alpinum var. pannonicum Terpo. and Arum alpinum var. intermedium Schur. was not found at Bilje (Tab. 5). Acknowlegement The authors would like to express their sincere gratitude to Professor Hazel Wilkinson, of the Botanic Gardens Kew, Jodrell Laboratory, for her great help in the stomata determination. References Baranova M. A. 1972; Systematic anatomy of the leaf epidermis in the Magnoliaceae and some related families. Taxon. 21 (4), 447-69. Bedalov M. 1981; Cytotaxonomy of the genus Arum (Araceae) in the Balkans and the Aegean area. Bot. Jahrb. Syst., pp. 183-200. Bedalov M. & Hesse M. 1989; On some pollen types within Arum (Araceae). Pl. Syst. Evol. 166: 41-44. 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I think, it should be introduced what mean the numbers in tables 1 - 3 Supplement Table 1 Stomata type and their prevalence at upper and lower epidermis in Arum italicum Mill., Arum maculatum var. maculatum L. and Arum maculatum var. immaculatum L. at location Zablae LocationZablaeEpidermisUpper epidermisLower epidermisType SHAPE \* MERGEFORMAT Arum italicum Mill.Arum maculatum var. maculatum L.Arum maculatum var. immaculatum L.Arum italicum Mill.Arum maculatum var. maculatum L.Arum maculatum var. immaculatum L.Paracytic4.503.433.973.404.804.33Hexacytic3.203.552.702.632.802.50Tetracytic1.751.831.401.100.630.63Brachyparacytic0.330.901.470.431.101.37Staurocytic0.500.200.350.370.250.73Anisocytic0.100.100.100.170.350.40Helicocytic0.000.000.000.100.000.00Anomocytic0.000.100.100.000.000.00Hemiparacytic0.100.100.000.100.000.00Hemidiacytic0.000.150.250.000.170.27 Table 2 Stomata type and their prevalence at upper and lower epidermis in Arum italicum Mill., Arum maculatum var. maculatum L. and Arum maculatum var. immaculatum L. at location Normanci LocationNormanciEpidermisUpper epidermisLower epidermisType SHAPE \* MERGEFORMAT Arum italicum Mill.Arum maculatum var. maculatum L.Arum maculatum var. immaculatum L.Arum italicum Mill.Arum maculatum var. maculatum L.Arum maculatum var. immaculatum L.Paracytic2.674.773.734.175.334.13Hexacytic3.132.203.032.672.002.70Tetracytic1.601.031.270.470.470.53Brachyparacytic2.001.731.531.631.431.73Staurocytic0.000.200.250.100.600.23Anisocytic0.100.100.000.100.100.10Helicocytic0.230.200.151.200.000.10Anomocytic0.100.100.000.000.100.00Hemiparacytic0.000.000.000.000.000.00Hemidiacytic0.300.100.173.330.170.53 Table 3 Stomata type and their prevalence at upper and lower epidermis in Arum alpinum var. pannonicum Terpo. and Arum alpinum var. intermedium Schur. at location Bilje LocationBiljeEpidermisUpper epidermisLower epidermisType SHAPE \* MERGEFORMAT Arum alpinum var. pannonicum Terpo.Arum alpinum var. intermedium Schur.Arum alpinum var. pannonicum Terpo.Arum alpinum var. intermedium Schur.Paracytic3.073.805.475.50Hexacytic3.472.802.501.77Tetracytic0.900.930.200.10Brachyparacytic1.772.001.371.67Staurocytic0.250.100.200.35Anisocytic0.100.000.000.25Helicocytic0.300.150.100.20Anomocytic0.000.100.100.00Hemiparacytic0.000.000.000.00Hemidiacytic0.470.200.271.93 Table 4 Mean number of stomata per square mm at upper and lower epidermis in some Arum species (t- test) at locations Zablae and Normanci LocationZablaeNormanciSpeciesArum italicum Mill.Arum maculatum var. maculatum L.Arum maculatum var. immaculatum L.Arum italicum Mill.Arum maculatum var. maculatum L.Arum maculatum var. immaculatum L.EpidermisUpperLowerUpperLowerUpperLowerUpperLowerUpperLowerUpperLower EMBED Equation.3 26.40235.0036.50196.0033.10290.0024.90651.0031.70305.0038.00403.00S.D.26.00111.0032.30163.0025.20147.0047.40287.00ace27.40117.0021.30220.00bd EMBED Equation.3  = mean value S.D. = standard deviation Table 5 Mean number of stomata per square mm at upper and lower epidermis in some Arum species (t- test) at locations Normanci and Bilje LocationNormanciBiljeSpeciesArum italicum Mill.Arum maculatum var. maculatum L.Arum maculatum var. immaculatum L.Arum alpinum var. pannonicum Terpo.Arum alpinum var. intermedium Schur.EpidermisUpperLowerUpperLowerUpperLowerUpperLowerUpperLower EMBED Equation.3 24,965131,73053840310639776,3381S.D.47,428727,4117a21,3220.00b9919254,1245  EMBED Equation.3  = mean value S.D. = standard deviation  Fig 1 Paracytic type in Arum italicum (BUWcˢТڢiN5ht8hOJQJcHdhdhdh^mH sH 5ht8h/3OJQJcHdhdhdhڙmH sH 5ht8hOJQJcHdhdhdh]mH sH #Hh]hOJQJmH sH ht8hOJQJmH sH ht85OJQJmHsHht8OJQJmHsHht85OJQJmH sH ht8:OJQJmH sH ht8OJQJmH sH DC$Eƀ] p#  p#$a$ " $$Ifa$  @$If$dha$$a$ "Ф*FRh &'()*+9?MQ÷zzfTzEEht86CJOJQJmHsH#j[ht8CJOJQJUmH sH &jht8CJOJQJUmHnHu#jht8CJOJQJUmH sH ht8CJOJQJmHsHht86CJOJQJmH sH ht8CJOJQJmH sH ht8OJQJmH sH 5ht8hOJQJcHdhdhdh^mH sH BHhꊪFht8hOJQJcHdhdhdh^mH sH ҥ yy $$Ifa$  @$Ifrkd$$IfTlT0B&04 laT +?`ۦysjjjjjj 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(Zablae-upper epidermis leaf top) Scale bar=10m  Fig 2 Hexacytic type in Arum italicum Mill. (Zablae-lower epidermis-leaf middle) Scale bar=10m  Fig 3 Tetracytic type in Arum italicum Mill. (Zablae-upper epidermis-leaf top) Scale bar=10m  Fig 4 Brachyparacytic type in Arum italicum Mill. (Zablae-upper epidermis-leaf top) Scale bar =10m   Fig 5 Helicocytic type In Arum italicum Mill. (Zablae-upper epidermis-leaf middle) Scale bar=10m  Fig 6 Anisocytic type in Arum italicum Mill. (Zablae-upper epidermis-leaf top) Scale bar=10m  Fig 7 Anomocytic type in Arum italicum Mill. (Zablae -lower epidermis-leaf middle) Scale bar=10m  Fig 8 Hemidiacytic type in Arum italicum Mill. (Zablae-lower epidermis-leaf top) Scale bar=10m  Fig 9 Hemiparacytic type in Arum italicum Mill. (Zablae-lower epidermis-leaf top) Scale bar=10m  Fig 10 Staurocytic type in Arum italicum Mill. 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