Peculiarities in the formation of the andrecium and gynecium elements are described for Arabidopsis plants grown from the stages of two cotyledonous leaves in the Svitoblok-1 device on board the Salyut 6 orbital research station and in the laboratory. It is established that flower buds and flowers, normally formed in habitus, contain sterile elements of andrecium and gynecium whose degeneration occurs at different developmental stages of the Arabidopsis plants in the experiment under conditions of weightlessness

Chernyayeva, I. I.

Kordyum, Y. L.

English

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NASA TECHNICAL MEMORANDUM NASA TM-77263
PECULIARITIES IN FORMATION OF ARABIDOPSIS THALIANA (L. ) HEYNH.
GENERATIVE ORGANS UNDER SPACE FLIGHT CONDITIONS
Ye. L. Kordyum and I. I. Chernyayeva
Translation of "Osoblyvosti formuvannya h;enerai-vvnykh orhaniv Arabidopsis
thaliana (L. ) Heynh. v umovakh kosmichnoho pol'yqhj," Dopovidi Akademiyi
nauk Ukrayinskoyi RSR, seriya B, no. 8, 198^, pp. 67-70.
NATIONAL AERONAUTICS AND SPACE ADMINISTRATION
WASHINGTON, D.C. 20546 JULY 1983
https://ntrs.nasa.gov/search.jsp?R=19830024006 2020-03-21T03:26:30+00:00Z
STANDARD TITLE PAGE
1. Rtport No.
NASA TM-77263
3. Govimatnt Acctltion No. >. RcclpUnl'* Cotoloj Na.
4. Till* mi SublilU
PECULIARITIES IN FORMATION OF ARABIDOP-
5. R*porl Dot*
July 1983
SIS THALIANA (L.) HEYNH. GENERATIVE
ORGANS UNDER SPACE FLIGHT CONDITIONS
6. Reforming Otgoniaotion Cod*
7. Aulhor(i)
Ye. L. Kordyum and I. I. Chernyayeva
Ukrainian SSR Academy* of Sciences
Botanical Institute
8. Pat farming Orgonizollan Report No.
10. Work Unit No.
9. Ptrforming Ofgcnuction Ncma and Add'dl
Leo Kanner Associates
Redwood City, CA 94063
II. Contract of Grant No.
NASW -3541
13. Type of Rtpo'l and Ptriod CovtrH
Translation
-12, Sponlorma Agincy Norn* end Addrtd
National Aeronautics and Space Adminis-
tration, Washington, D.C. 20546 14. Sponsoring Agency Cod*
15. SvppUmtntcry Not**
Translation of "Osoblyvost-i formuvannya henerativnykh orhaniv
Arabidopsis thaliana (L.) Heynh. v umovakh kosmichnoho
pol'yotu," Dopovidi Akademiyi nauk. Ukrayinskoyi RSR, seriya B
'no. 8, 1982, pp. 67-70.
16. Abstto'ci
Peculiarities in the formation of the andrecium and gynecium
elements are described for Arabidopsis plants grown from the
stages of two cotyledonous leaves in the.Svitoblok-1 device
on board, the Salyu't 6 orbital research station and in the
laboratory. It is- established that flower buds and flowers,
normally formed in habitus, contain sterile elements of
andrec'ium and gynecium whose degener'ation occurs at different
developmental stages of the Arabidopsis plants in the
experiment under cpnditions of weightlessness.
17. Ktj Word* (Selected by Author(i)} 18. Distribution StoUmtnl
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"• FROM
PECULIARITIES IN FORMATION OF ARABIDOPSIS THALIANA (L.) HEYNH.
GENERATIVE ORGANS UNDER SPACE FLIGHT CONDITIONS
Ye. L. Kordyum and I. I. Chernyayeva
Ukrainian SSR Academy of Sciences Botanical Institute
The possibility of using annual higher plants in complete ontogene-/67*
sis (from seed to seed) under space flight conditions has been examined
in some articles [1-5], but there has been no experimental, confirmation.
Aboard the Salyut-6 orbiting scientific space station an experiment
was done which involved growing Arabidopsis, an ephemeral of the family
Cruciferae (Brassicaceae)—a convenient subject for radiobiological and
space biological experimentation [6]—in a "Svitloblok-1" apparatus,
which permitted growth of small higher plants under sterile, sealed
conditions on an agarized growth medium, with light from 2 to 4 thousand
lux . Arabidopsis seedlings in the dicotyledonous phase were placed
in the apparatus and put aboard the Salyut-6. The experiment lasted 65
days. For the first 16 days the plants were illuminated 14 hrs a "day,
for the last 51 days they were illuminated around-the-clock. The con-
trol plants were grown under laboratory conditions with the same
lighting schedule.
At the end of the experiment, both the control and experimental
plants were in the same stage of growth — blossom and, spadix fertil-
ized — and were no different as to average stem length, number of buds,
flowers, and pods. The low light intensity produced a slowing of the
generative phase. Arabidopsis is a long-day plant, its ontogenesis
accomplished in 45-47 days if light intensity is 10 thousand lux and
day length is no less than 16 hours.
For examination with the light microscope, buds, flowers, and pods
of the Arabidopsis controls and experimental plants were fixed in 2.5%
*Numbers-in the margin indicate pagination in the foreign text.
1. Seeds produced at the USSR Academy of Sciences Institute of
General Genetics.
glutaraldehyde, with postfixation in a 1% solution of Os04 and Navashin's
mixture (chromacetoformol). The material was dried and embedded in /68
paraffin using the accepted cytologic method. Sections 10 ym thick
were obtained with an MPS-2 microtome, stained with Heidenhain's
iron hematoxylin and methyl green, and examined under an "NF" micro-,
scope. Sections were also stained using Feulgen's and Lugol's reactions
and Sudan III; the latter two staining methods were used to ascertain
the nature of nutrient stores in the pollen grain vegetative cells.
The preparations were sketched using an RA-4 drawing apparatus and
photographed with an "NF" photographic attachment.
Microscopic examination of the Arabidopsis plant generative organs
in the last stages of formation, as grown in the "Svitoblok-1" appar-
atus under laboratory conditions, revealed that the formation and
development of gynecium and andrecium elements proceeded normally
(see figure, A), as described in [7-8].
It was established, however, that the plants in the experimental
population were not obligate self-pollinating, since, in addition to
the normally first-order hermaphroditic flowers, potentially bisexual
female ones had^fdrmed;- These flowers differed from the hermaphroditic
ones in external appearance. Before blossoming of the hermaphroditic
ones, sepal length was no longer than 2/3 that of the white petals,
which remained closed as the pollen burst out, ensuring contact between
the pollen niduses of the long stamens, which had already opened, and
the stigma, thus allowing self-pollination. In the potentially bisexual
female flowers the petals did not exceed or only somewhat exceeded the
sepals, so that the perianth surrounded no more than 2/3 the length
of the ovule (the germ of which is seen on top), due to which cross-
-pollination is possible under natural conditions. When these flowers
were artificially pollinated with the pollen of other plants, pods
formed with complete seeds. The plants grown from these seeds were
fertile. Thus this population of Arabidopsis was of gynomonoecious
sexual type.
As opposed to the control plants, whose green pods contained seeds
with germs at various stages of development, including differentiated,
the experimental plants were found to have parthenocarpic pods with
degenerating seed embryos. The buds and seeds apparently normally .
formed were replaced by sterile andrecium and gynecium elements whose
degeneration began at various stages of generative organ development
(see figure). It is remarkable that, as a rule, the flowers in the
experimental material were externally similar to the potentially
bisexual female flowers in the controls, i.e., the stigma and style
and, generally, part of the germ were not surrounded by perianth,
indicating the presence of a positive correlation between the degree
of andrecium fertility and stamen filament length and size and location
of the petal panicle, i.e., a morphological mechanism for ensuring
pollination — either self- or cross-pollination.
Andrecium element degeneration in the experimental plants began
at the tetrad microspore or mononuclear microspore stages. Niduses
, rich in pollen were found to have bicellular or even tricellular pollen
grains, but appeared non-viable "from outward signs:— the absence in the
vegetative cell cytoplasm of sufficient nutrient reserves, particularly
of fat. The microsporangium walls, as in the controls, were made up of
four layers of cells: epidermis, endothecium — the future fibrous
layer, middle layer, and secretory-type tapetum with binuclear cells;
in the mature state -- epidermis and fibrous layer.
Gynecium element degeneration was found at the tetrad microspore,
mono-, bi-, and tetranuclear stages, and in the fomed embryonal sac.
In none of the buds studied did the establishment or first developmental
stages of the seed embryos differ from those in the controls. Two
clear pathways of sterilization were later seen -- synchronous cessation /70
of macrospore or seed embryo embryonal sac and somatic cell development,
and asynchronicity of embryonal sac and somatic element (primarily
integument) element development, the first signs of developmental ces-
sation appearing, as a rule, in the somatic elements. These primarily
were the absence of integumentary tapetum differentiation, partial lysis
of internal integument cells, and pathological vacuolization of integ-
ument cells. At the same time, the enriched seed embryos had morpho-
3
Arabidopsis thaliana (L.) Heynh.
A — Control: a — appearance of hermaphroditic flower in
longitudinal section; b-e — fragments of a; b — tricellu-
lar pollen grain; c,d — seed embryos from mature embryonal
sacs; e — ovum; B — Experimental: a -- appearance of flower
in longitudinal section; b-d — fragments of a; b,c — sterile
microspores; d -- sterile seed embryo.
logically normally formed embryonal sacs with ov (egg cell and two
synergids) and secondary nucleus of embryonal sac central cell. The
mature seed embryos had a more or less campylotropous form, character-
istic of seed embryos of this species, but they were smaller than in
the controls. No fertile seed embryos were formed.
Thus the morphogenetic processes in the generative organs were
more or less the same under weightless conditions, within the parameters
used in this experiment, as normal. Generative organ sterilization
was caused, most likely, by pathologically altered processes at the
cell level, primarily cessation of cell division and differentiation,
which have their specific structural manifestations in the andrecium
and gynecium.
Based on comparative analysis of sterilization process features in
the andrecium and gynecium elements in the experimental Arabidopsis
plants and other angiospermous plants with various types of sterility
under natural conditions, as well as possible causes of nondifferenti-
ation of integumentary tapetum in seed embryos [9], it is assumed that
the natural hormonal nature was disrupted in the generative phase of
Arabidopsis plant development in this experiment under weightless
conditions.
REFERENCES
1. Merkys, A. J., A. L. Mashinsky, R. S. Lauriavickus et al. , "The
development of seedling shoots under space flight conditions,"
Life Sci. and Space Res. 13, 53-57 (1975).
2. Merkis, A. I., "The importance of gravity in plant growth processes,"
Probl. kosm. biologii 33, 146-174 (1976).
3. Platonova, R. N., G. P. Parfenov, V. P. Ol'khovenko et al., "Pine
seed growth in weightlessness (study on Cosmos-782 satellite),"
Izv. AN SSSR, Ser. biologiya, 5, 770-776 (1977).
4. Sitnik, K. M., and Ye. L. Kordyunt, "Botanical research in space,"
Ukr. hot, zhurn. 37/1, 1-10 (1980).
5. Parfenov, G. P., and V. M. Abramova, "Blossoming and maturation of
Arabidopsis plants in weightlessness (experiment on Cosmos-1129
biosatellite)," Dokl. AN SSSR 256/1, 254-256 (1981).
6. Ivanov, V. I., "Radiobiology and genetics of Arabidopsis," Probl.
kosm. biologii 27, 38-50 (1974).
7. Polyakova, T. F., "Development of male and female gametophytes of
Arabidopsis thaliana (L.)," Issled. po genetike, 2, 125-133 (1964).
8. Yakovlev, M. S., and G. K. Alimova, "Embryogenesis in Arabidopsis
thaliana (L.) Heynh. (Cruciferae) , u Botan. zhurn. 61/1, 12-24
(1976).
9. Kapil, R. N., and S. C. Tiwari, "The integumentary tapetum," Bot.
Rev. 44/4, 457-490 (1978).


Peculiarities in formation of Arabidopsis Thaliana (L.) Heynh, generative organs under space flight conditions

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