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Przypisy oksfordzkie w treści głównej są linkami oznaczonymi słowem „footnote” i numerem porządkowym wziętymi w nawias kwadratowy, a przypisy harvardzkie słowem „footnote”, numerem porządkowym, nazwiskami autorów oraz rokiem wydania wziętymi w nawias okrą

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Uwagi do wersji zaadaptowanej:

Wersja elektroniczna książki została stworzona zgodnie z art. 33 z indeksem 1 Ustawy o prawie autorskim i prawach pokrewnych.

Zostały zachowane numery stron. Numer danej strony znajduje się nad tekstem danej strony i poprzedza go skrót str.

Wartości wyrażone w oryginale liczbami rzymskimi w adaptacji przedstawiono cyframi arabskimi albo słownie.

Przypisy oksfordzkie w treści głównej są linkami oznaczonymi słowem „footnote” i

numerem porządkowym wziętymi w nawias kwadratowy, a przypisy harvardzkie słowem

„footnote”, numerem porządkowym, nazwiskami autorów oraz rokiem wydania wziętymi w nawias okrągły. Linki te prowadzą do opisów przypisów, które zostały umieszczone na końcu dokumentu. Powrót do treści głównej po odczytaniu opisu przypisu jest możliwy poprzez link „Return to the main document” umieszczony na końcu każdego opisu przypisu.

Zakładki oznaczone literą „f.” i „ff.” oraz numerem porządkowym znajdują się w treści głównej, w miejscu występowania przypisu.

Wykaz skrótów:

cv. – cultivar str. – street var. – various

Desf. – nazwa gatunku, której autorem jest René Louiche Desfontaines DNA – deoxyribonucleic acid

L – nazwa gatunku, której autorem jest Karol Linneusz Mill – nazwa gatunku opisana przez Phillipa Millera PCD – programmed cell death

SEM – Scanning electron microscopy Koniec uwag do wersji zaadaptowanej.

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Modern Phytomorphology 6: 19-20, 2014

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ISOLATED ENDOSPERM TISSUE AS A MODEL FOR EXPERIMENTAL BOTANY MARZENA POPIELARSKA-KONIECZNA [footnot e 1 ], MAŁGORZATA KOZIERADZKA- KISZKURNO [footn ote 2 ], IZABELA MARCIŃSKA [footnote 3 ], DAGMARA KWOLEK [footnote 4 ], HALINA ŚLESAK [footnote 5 ]

Abstract. Experiments conducted on isolated endosperm under in vitro conditions have opened new possibilities to investigate this specific and unique plant tissue. Results are important as well for basic knowledge as for agriculture practice.

Key words: cereals, kiwifruit, regeneration, triploid plants

Endosperm is the ephemeral tissue, however it plays an important role in the life cycle of angiosperms plant. In most angiosperms, endosperm is formed during the process of double fertilization as a product of the fusion of a haploid sperm nucleus with two haploid polar nuclei. It results in a triploid structure formation, which develops into the tissue extreme specialized (footnote 1. Costa L.M., Gutièrrez-Marcos J.F., Dickinson H.G. 2004 ) and consumed by embryo during embryogenesis or seed germination. Interesting, there are no reports concerning in vivo endosperm differentiation resulted in plant regeneration.

But it was proved that endosperm under in vitro cultures has the potential to proliferate, differentiate and finally even plant regenerate.

Both, mature and immature endosperm tissues revealed ability to proliferate in vitro.

Factors like proper stage of endosperm development and culture conditions (e.g. plant growth regulators, light conditions) are important for successful plant regeneration. The capacity for tissue proliferation and morphogenetic response differ among species. There could be observed the correlation between nutrition status of plant and a kind of

morphogenetic reaction. Usually, direct organogenesis is typical for parasitic and

semiparasitic species. The semi-parasitic Exocarpus cupressiformis Labill. was indeed the first species ever to show the totipotency of endosperm (footnote 2. Johri B.M. & Bhojwani S.S. 1965 ). The endosperm of autotrophic plant indicated indirect organogenesis and callus stage is necessary for next steps of differentiation. Continuously growing non- morphogenic callus cultures, without organ regeneration, have been obtained from the endosperm of such important commercial species like maize (Zea mays L.), cocos (Cocos nucifera L.), tomato (Lycopersicon esculentum Mill.) or cucumber (Cucumis sativus L.) (footnote 3. Thomas T.D. & Chaturvedi R. 2008). Plants regenerated this way show very often 3C level of nuclear DNA content. Successful 3C plant regeneration from endosperm-

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derived callus has been reported actually in 19 species, e.g. in acacia (Acacia nilotica (L.) Delile), walnut (Juglans regia L.), papaya (Caricapapaya L.).

Kiwifruit (Actinidia deliciosa (Auguste Chevalier) C.F. Liang et A.R. Ferguson var.

deliciosa) is one of the important crops in interests of our team. We described efficient protocol for plant regeneration from endosperm-derived callus (footnote 4. Góralski G., Popielarska M., Ślesak H., Siwińska D., Batycka M. 2005). Kiwifruit is hexaploid (2n equals 6x equals174) and because of that flow cytometry was used to determine the ploidy of callus and regenerated organs. We investigated the differences (e.g. the presence of the plant extracellular matrix, cutin, pectins) between morphogenic and non-morphogenic endosperm-derived callus in kiwifruit (footnote 5. Popielarska M., Góralski G., Ślesak H.

2006; Popielarska-Konieczna M., Kozieradzka - Kiszkurno M., Świerczyńska J., Góralski G., Ślesak H., Bohdanowicz J. 2008, Popielarska-Konieczna M., Kozieradzka - Kiszkurno M., Bohdanowicz J. 2011).

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The other object of our research is cereals. Its isolated immature endosperm develops under in vitro conditions in similar way like in caryopsis in planta. Recently (footnote 6.

Popielarska-Konieczna M., Kozieradzka - Kiszkurno M., Tuleja M., Ślesak H., Kapusta P., Marcińska I., Bohdanowicz J. 2013) we conducted detailed histological and ultrastructural studies on isolated endosperm of bread wheat (Triticum aestivum L.), durum wheat

(Triticum durum Desf.) and triticale (Triticosecale Wittmack). Endosperm development of cereals is strongly connected with the starch accumulation and programmed cell death (PCD) (footnote 7. Sabelli P.A. & Larkins B.A. 2009). In triticale induction of PCD starts in 16 days post anthesis (footnote 8. Li C.-Y., Li W.-H., Li C., Gaudet D.A., Laroche A., Cao L.-P., Lu Z.-X. 2010). Our data (not published) revealed that cultured endosperm tissue of triticale, which accumulated starch granules under in vitro conditions showed the viability during 3-4 months of the culture, what was confirmed using Evans blue staining procedure (according to footnote 9. Zhou Z., Wang L., Li J., Song X., Yang C. 2009).

Recent reports (footnote 10. Carciofi M., Blennow A., Nielsen M.M., Holm P.B., Hebelstrup K.H. 2012) pointed that experimental research concerning endosperm-tissue are still needed and could be convenient platform to manipulation of the development and study the molecular and biochemical mechanisms regulating seed storage accumulation in the endosperm.

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Acknowledgments

The present works in part are financially supported by grant number 2012/07/b/

NZ9/01325 from The National Science Centre (Poland).

References

Carciofi M., Blennow A., Nielsen M.M., Holm P.B., Hebelstrup K.H. 2012. Barley callus:

a model system for bioengineering of starch in cereals. Plant Methods 8: 36.

Costa L.M., Gutièrrez-Marcos J.F., Dickinson H.G. 2004. More than a yolk: the short life and complex times of the plant endosperm. Trends Plant Science 9: 507-514.

Góralski G., Popielarska M., Ślesak H., Siwińska D., Batycka M. 2005. Organogenesis in endosperm of Actinidia deliciosa cv. Hayward cultured in vitro. Acta Biologica

Cracoviensia Series Botanica 47: 121-128.

Johri B.M., Bhojwani S.S. 1965. Growth response of mature endosperm in cultures.

Nature 298: 1345-1347.

Li C.-Y., Li W.-H., Li C., Gaudet D.A., Laroche A., Cao L.-P., Lu Z.-X. 2010. Starch synthesis and programmed cell death during endosperm development in triticale (x Triticosecale Wittmack). J. Integrative Plant Biology 52: 602-615.

Li J., Berger F. 2012. Endosperm: food for humankind and fodder for scientific discoveries. New Phytologist 195: 290-305.

Popielarska M., Góralski G., Ślesak H. 2006. Histological and SEM studies on

organogenesis in endosperm-derived callus of kiwifruit (Actinidia deliciosa cv. Hayward).

Acta Biologica Cracoviensia Series Botanica 48: 97-104.

Popielarska-Konieczna M., Kozieradzka-Kiszkurno M., Świerczyńska J., Góralski G., Ślesak H., Bohdanowicz J. 2008. Ultrastructure and histochemical analysis of

extracellular matrix surface network in kiwifruit endosperm-derived callus culture. Plant Cell Reports 27: 1137-1145.

Popielarska-Konieczna M., Kozieradzka-Kiszkurno M., Bohdanowicz J. 2011. Cutin play a role in differentiation of endosperm-derived callus of kiwifruit. Plant Cell Reports 30: 2143- 2152.

Popielarska-Konieczna M., Kozieradzka-Kiszkurno M., Tuleja M., Ślesak H., Kapusta P., Marcińska I., Bohdanowicz J. 2013. Genotype-dependent efficiency of endosperm development in culture of selected cereals: histological and ultrastructural studies.

Protoplasma 250: 361-369.

Sabelli P.A., Larkins B.A. 2009. The development of endosperm in grasses. Plant Physiology 149: 14-26.

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Thomas T.D., Chaturvedi R. 2008. Endosperm culture: a novel method for triploid plant reproduction. Plant Cell Tissue Organ Culture 93:1-14.

Zhou Z., Wang L., Li J., Song X., Yang C. 2009. Study on programmed cell death and dynamic changes of starch accumulation in pericarp cells of Triticum aestivum L.

Protoplasma 236: 49-58.

Footnotes

Footnote 1. Department of Plant Cytology and Embryology, Jagiellonian University,

Gronostajowa str. 8, 30-387 Cracow, Poland; m.popielarska-konieczna@uj.edu.pl. Return to the main document.

Footnote 2. Department of Plant Cytology and Embryology, University of Gdańsk, Wita Stwosza str. 59, 80-308 Gdańsk, Poland. Return to the main document.

Footnote 3. Institute of Plant Physiology of Polish Academy of Sciences, Niezapominajek str. 21, 30-239 Cracow, Poland. Return to the main document.

Footnote 4. Department of Plant Cytology and Embryology, Jagiellonian University, Gronostajowa str. 8, 30-387 Cracow, Poland. Return to the main document.

Footnote 5. Department of Plant Cytology and Embryology, Jagiellonian University, Gronostajowa str. 8, 30-387 Cracow, Poland. Return to the main document.

Footnotes (harvard)

Footnote 1. Costa L.M., Gutièrrez-Marcos J.F., Dickinson H.G. 2004. More than a yolk: the short life and complex times of the plant endosperm. Trends Plant Science 9: 507–514.

Return to the main document.

Footnote 2. Johri B.M., Bhojwani S.S. 1965. Growth response of mature endosperm in cultures. Nature 298: 1345–1347. Return to the main document.

Footnote 3. Thomas T.D., Chaturvedi R. 2008. Endosperm culture: a novel method for triploid plant reproduction. Plant Cell Tissue. Organ Culture 93:1–14. Return to the main document.

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Footnote 4. Góralski G., Popielarska M., Ślesak H., Siwińska D., Batycka M. 2005.

Organogenesis in endosperm of Actinidia deliciosa cv. Hayward cultured in vitro. Acta Biologica Cracoviensia Series Botanica 47: 121–128. Return to the main document.

Footnote 5. Popielarska M., Góralski G., Ślesak H. 2006. Histological and SEM studies on organogenesis in endosperm-derived callus of kiwifruit (Actinidia deliciosa cv. Hayward).

Acta Biologica Cracoviensia Series Botanica. 48: 97–104; Popielarska-Konieczna M., Kozieradzka-Kiszkurno M., Świerczyńska J., Góralski G., Ślesak H., Bohdanowicz J. 2008.

Ultrastructure and histochemical analysis of extracellular matrix surface network in kiwifruit endosperm-derived callus culture. Plant Cell Reports 27: 1137–1145; Popielarska-

Konieczna M., Kozieradzka-Kiszkurno M., Bohdanowicz J. 2011. Cutin play a role in differentiation of endosperm-derived callus of kiwifruit. Plant Cell Reports 30: 2143–2152.

Return to the main document.

Footnote 6. Popielarska-Konieczna M., Kozieradzka-Kiszkurno M., Tuleja M., Ślesak H., Kapusta P., Marcińska I., Bohdanowicz J. 2013. Genotype-dependent efficiency of endosperm development in culture of selected cereals: histological and ultrastructural studies. Protoplasma 250: 361–369. Return to the main document.

Footnote 7. Sabelli P.A., Larkins B.A. 2009. The development of endosperm in grasses.

Plant Physiology 149: 14–26. Return to the main document.

Footnote 8. Li C.-Y., Li W.-H., Li C., Gaudet D.A., Laroche A., Cao L.-P., Lu Z.-X. 2010.

Starch synthesis and programmed cell death during endosperm development in triticale (x Triticosecale Wittmack). J. Integrative Plant Biologica 52: 602–615. Return to the main document.

Footnote 9. Zhou Z., Wang L., Li J., Song X., Yang C. 2009. Study on programmed cell death and dynamic changes of starch accumulation in pericarp cells of Triticum aestivum L. Protoplasma 236: 49–58. Return to the main document.

Footnote 10. Carciofi M., Blennow A., Nielsen M.M., Holm P.B., Hebelstrup K.H. 2012.

Barley callus: a model system for bioengineering of starch in cereals. Plant Methods 8: 36.

Return to the main document.

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