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4
RESEARCH PAPER
RESEARCH PAPERRESEARCH PAPER
RESEARCH PAPER
OPEN ACCESS
OPEN ACCESSOPEN ACCESS
OPEN ACCESS
Biology o lowe ing and emb yological s uc u e o he i on
ee in ade onic condi ions
Gulna Ch.
Gasimo a
1
, Panah Z. Mu ado
*2
1
Ins i u e o Bo any, Minis y o Science and Educa ion, Republic o Aze baijan
2Ins i u e o Mic obiologyu, Minis y o Science and Educa ion, Republic o Aze baijan
Key wo ds:
Pa o ia pe sica, Rep oduc i e o gans, Seed o ma ion, Emb yological s udies
h p://dx.doi.o g/10.12692/ijb/25.2.10-16 A icle published on Augus 03, 2024
Abs ac
The a icle s udies he de elopmen o he ep oduc i e o gans o i on
ee
, which belongs o he elic lo a o
he hi d pe iod. I has been es ablished ha Pa o ia pe sica
has no se ious obs acles ha impede he no mal
p ocess o seed o ma ion. The lowe g oup o his succulen species is almos de oid o co e . Mo eo e , i is
mainly o med du ing he win e season,
which limi s he in oduc ion o he plan in o mo e se e e clima ic
zones. İn he Abshe on Peninsula, i on ees bloom and bea ui only in yea s wi h mild win e s, s ong
winds and he absence o nigh
os s. Emb yological s udies ha e es ablished co espondence be ween he
unc ions o he epide mis and endo hecium, he p esence in males o a clea ly isible al e, in which he
nuclei o he ape al cells a e loca ed linea ly in he adial di ec ion, as
well as a delay in he de elopmen o
he emale geni al o gans compa ed o he male sys em.
*
Co esponding Au ho : Panah Z. Mu ado mpanah@mail. u
In e na ional Jou nal o Biosciences | IJB |
ISSN: 2220-6655 (P in ) 2222-
5234 (Online)
h p://www
.innspub.ne
Vol. 25, No. 2, p. 10-16, 202
4
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4
In oduc ion
Pa o ia pe sica (D.C.) C.A.-a unique bo anical
objec wi h o iginal biological p ope ies. Maybe
oday i has become he cen e o a en ion o
esea che s. A numbe o aspec s o he de elopmen
p ocess o his elic plan , which is one o he li ing
elic s o he Te ia y pe iod, including i s
emb yology, ha e been p ac ically no s udied.
Pa o ia pe sica, conside ed one o he aluable
woody plan s, is na u ally dis ibu ed only in he
Hy сan o es s loca ed in he Lanka an egion o
Aze baijan, and in he local o es s o he Gabala
egion. Due o he high speci ic g a i y o his plan ,
i easily sinks in wa e , which is why Pa o ia pe sica
is popula ly called he “i on ee”. Obse a ions show
ha shoo s, lowe s and lea es o i on ee can
emain on he main plan o a long ime.
I should be no ed ha he s udy o he in icacies
o biology and emb yology o he p ocess o
o ma ion and lowe ing o he i on ee is s ill no
a a sa is ac o y le el. The e o e, s udying Pa o ia
pe sica in he abo e aspec s will u he inc ease
in e es in he in oduc ion and conse a ion o
i on ee. Also o g ea impo ance in sol ing he
p oblem is he s udy o he cha ac e is ics o he
de elopmen o shoo s in he i on ee, as well as
he pa e ns o o ma ion o he ep oduc i e
o gans. I is on he basis o he esul s ob ained in
he cou se o esea ch ca ied ou in his di ec ion
ha i is possible o conduc a compa a i e
emb yology o indi idual ep esen a i es o he
amily Hamamelidaceae. In o he wo ds, i is
possible o ind ou wha changes occu ed in he
elemen s o he ep oduc i e s uc u e du ing he
adap a ion o his ancien plan o mode n
en i onmen al condi ions along he pa h o
mo phological e olu ion.
Mono ypic genus Pa o ia pe sica (D.C.) C.A. Mey
belongs o he Hamamelidaceae amily.
Rep esen a i es o his amily we e widesp ead in
he hi d pe iod. Conduc ed paleon ological s udies
o his genus Pa o ia agi olia (Goepp.) Hee and
P. p is ina (E ingsh.) S u . shows ha i s species
we e once widesp ead. Pa o iopsis
jasquemon iana Dens, ound in he Himalayas
(moun ains o Kashmi ). The species a e e y
simila o membe s o he genus Pa o ia in all
hei biological cha ac e is ics.
Rep esen a i es o he Hamamelidaceae amily a e
mo e common in opical and sub opical clima e
zones. These plan s a e no ound in a eas wi h
ad e se na u al condi ions. The i s s udies ela ed o
he s udy o he biological p ope ies o i on ee in
ou epublic we e ca ied ou by Sa a o in 1960-1977
yea s. Thus, he conduc ed scien i ic esea ch on he
p opaga ion o his plan by seeds, biology, ecology
and phy ogenology o i s lowe ing, and i s cul i a ion
unde cul u al condi ions (Sa a o , 1972; 1977). In
addi ion, al hough weakly, bu also s udied mycobio a
o his ee (Mu ado e al., 2020). Howe e ,
emb yological s udies, especially he de elopmen o
ep oduc i e o gans o he i on ee a e p ac ically no
ca ied ou , al hough such s udies (Janssens e al.,
2020, Rensing and Weije s, 2021) a e impo an o
unde s anding he gene al de elopmen o his ee.
In his ega d, he pu pose o he p esen ed wo k
was o s udy he de elopmen o ep oduc i e o gans
in he condi ions o Aze baijan.
Ma e ials and me hods
The ma e ial o mo phological analysis and
emb yological esea ch was aken om 30-35-yea -
old Pa o ia pe sica ees g owing in he Cen al
Bo anical Ga den. To p epa e s able p epa a ions,
he ma e ial was ixed in Ca noy's solu ion (6:3:1).
Emb yological ma e ial was p epa ed using he
gene ally accep ed cy ological (Paushe a, 1974). To
s ain s able p epa a ions, hey a e s ained wi h
bo h hemo oxin and Alcian blue solu ion acco ding
o Kede enghain. Pollen e ili y was s udied by
s aining wi h ace oca mine; iabili y was
de e mined du ing ge mina ion in a 10-15% suga
solu ion by he d op me hod. Pollen mo phology
was s udied using a uchsin s ain. S abilized
p epa a ions we e s udied and pho og aphed using
MBI-6 and ZEISS S emi 2000-C mic oscopes,
Oxion eu omex.
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In . J. Biosci.
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4
Resul s and discussion
Shoo s
As a esul o he s udies, i was es ablished ha he
shoo s o med by i on ee a e ei he ege a i e o
mixed in na u e. A he end o au umn,
mo phologically iden ical shoo s a e b own and ha e
sho legs. Top shoo s 4-5 mm, la e al shoo s 2-3 mm.
In au umn hey become e y ha d and consis o 1-3
udimen a y lea es. Each o hem p oduces wo wild
co ns in a bud. Obse a ions show ha he i on ee
does no ha e ue shoo lea es, and hei unc ion is
pe o med by pseudopods. To p o ec o e win e ing
emb yonic lea es and a ce ain pa o he
hododend on om un a o able en i onmen al
condi ions, 10 o mo e da k b own delica e hai s a e
o med (Fig. 1, i1). No e he appea ance o each hai
unde he mic oscope, he shape o an elonga ed
mononuclea cell. The hai s a e co e ed only by he
lea es o ege a i e shoo s (Fig. 1, and 2). Howe e ,
he ou e su ace o he lowe bed and o a y is
su ounded by cilia (Fig. 1, e; 3, a-d). Obse a ions
show ha he ac i e de elopmen o mixed and
ege a i e shoo s o i on ee ne e begins
simul aneously.
Fig. 1.
Mo phological de elopmen a di e en s ages o lowe and shoo
a- young b anches (a1) and mixed shoo s (a2) in Decembe ; b- Janua y- Feb ua y; -
ege a i e shoo s (b1) and
mixed shoo s (b2) (Ma ch); q- opened lowe s; d-
mixed shoo s, wo imma u e lea es in he axil o he lowe (2
lea es pe lea , he ips o he shoo s a e de o med); e- c oss sec ion o a lowe (pe iole 1; o lowe b
ed 6;
s amens 11; pe iole 2); j- young o a y; z- ui s; u1- b is les; u2- s ages o hai de elopmen
Mixed shoo ing
These shoo s o m on he mo he plan a he end o
No embe . A his ime, mixed shoo s (Fig. 1a; 1a2)
ake on a ounded shape, and ege a i e shoo s in he
sp ing a e no qui e simila o each o he (Fig. 1a;
1a2). In Janua y, he diame e o mixed shoo s
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In . J. Biosci.
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4
eaches 1 cm. Mixed shoo s consis s o one o wo
co yledons, on each o which he heads o la e al
lowe buds a e also isible. F om which 5-7 lowe -
emb yos a e o med. Du ing he lowe ing o i on o e,
he lowe c own o med om one lowe bed is b igh
g een o da k b own on he inside (Fig. 1b; 1g; 1d),
ba e and d ooping on he ou side.
The numbe o co ollas is equal o he numbe o
lowe s o med du ing lowe ing. C owns a e o med
on peonies wi h a diame e o 7 mm in he
lowe bed. La ge ones open in mid-Janua y and 30-
40 ed s amens appea . A his ime, he ounda ion
o o he elemen s o he lowe is laid. Howe e ,
hei isual image has no ye been eco ded.
The e o e, be o e lowe ing in Janua y, i on peonies
exhibi ba e and shiny s amens o a unique
mo phology. I should be no ed ha a his ime he
pollen is only in he uninuclea e s age, and he basis
o he seeds in he o a y has no ye been laid.
Peonies o e win e in i on ee in a semi-open s a e.
I on ee lowe s g own in na u al condi ions a e
p o ec ed in his s a e by lea es emaining on he
mo he plan om he p e ious yea . In oduced
i onwood in Abshe on condi ions is a ec ed by
s ong winds; lowe de elopmen is delayed due o
he ac ha all he lea es emaining om he
p e ious yea all o . Despi e he ac ha Pa o ia
pe sica is poo ly p o ec ed om un a o able win e
condi ions, i s ba e peonies a e ac i ely de eloping,
and his adap i e ea u e can be conside ed an
a a ism. I should be no ed ha his plan g ows
well in wa m wea he unde na u al condi ions. In
he wo ks o Sa a o (1977) p o ides in o ma ion
abou he exis ence o e e g een species o i on o e
and o he elic plan s o he hi d pe iod. Flowe
de elopmen in peonies begins almos
synch onously. The numbe o lowe s a anged
cyclically a ound he lowe bed a ies om 5-7. The
numbe o s amens in a lowe bed is uns able,
luc ua ing be ween 9-11 and hey a e loca ed
opposi e each o he .
Fig.
2
.
Fo ma ion o he male ep oduc i e s uc u e
a- ini ial de ec ion o he s amen in he an he , b- c oss-sec ion o he an he , - ans e sely cu an he s; -
an he wall in he meiosi is s age (Decembe ), q- diakinesis, d- me aphase, e- elophase I, j- p ophase II, z-
mic ospo e in a e ad, u- seed ge mina ing om he an he (end o Feb ua y), s ained wi h magen a; en, kl-
cellula spo ogens, o . old.- middle laye ; ind- ape um, end.- endo hecium, epi.- epide mis.
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In . J. Biosci.
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4
Fo ma ion o pollen sacs and an he s (Fig. 2a-2u).
The gene a ion and di e en ia ion o he and oecium
begins in la e No embe - ea ly Decembe . Young
pollen sacs, eaching a heigh o 1.5 mm in mid-
Decembe , soon begin o u n ed-pink and end wi h
meiosis. Du ing his pe iod, he wall o he pollen sac
becomes 6-7-laye ed, and he cell wall has a hickened
epide mis.
The cells o he subepide mal laye , i.e., endo hecin,
unlike o he angiospe ms, lack hickened ib osis.
These cells inc ease in size a di e en s ages o
pollen sac o ma ion and o en unde go changes. In
his case, hese cells unde go lysis along wi h he
middle laye and ape um. Cha ac e is ics o he
mo phological ea u es o he endo hecium and
epide mal cells show ha he speci ic unc ions o he
endo hecium a e pe o med by he epide mis. The
middle laye is 4, and he inne mos laye is la ened.
The sec e o y ape um is homogeneous. The cells
aken he e had 1–4 nuclei and we e highly elonga ed
in he angen ial di ec ion. In ape al cells, he
loca ion o he nuclei on he in e nal longi udinal axis
is o g ea in e es . The wid h o he ape al laye is
equal o he wid h o 3-4 middle laye s. In
mic ospo ocy es, meiosis usually occu s
synch onously. A his ime, he numbe o haploid
ch omosomes is n=12, and mic ospo es a e loca ed
e ahed ally. Thus, al hough he cha ac e is ics
men ioned abo e, including he mul i-laye ed pollen
sac, he hickness o he cell walls o he epide mis,
he s ong connec ion be ween he uppe and lowe
pa s o he beak-shaped s amen, seem p imi i e om
an e olu iona y poin o iew, hey a e ex emely
impo an in he p o ec ion o i onwood pollen, which
is a li ing elic his hi d pe iod, no only om hea , bu
also om cold.
In ea ly Janua y, young mic ospo es eme ge om he
shell in he o m o e ads. A he end o Feb ua y, wo-
celled e icula ed seeds co e ed wi h e icula e exine a e
o med. Ripe e icula ed i on o e seeds a e loca ed in he
me idional di ec ion wi h h ee g oo es and ha e a
ound shape wi h a diame e o 46 mic ons.
On Abshe on, in wa m au umn and windless win e ,
he e ili y o i onwood pollen is 90-92%; in a yea o
ho au umn and cold win e (-2; -4 C), he bi h a e
d ops o 40% o less. In some cold yea s, i onweed
only ege a es and does no bloom ( his happened in
2017). The dus pa icles c acked longi udinally.
Fi s , pollen nes s bloom in he middle pa along he
en i e leng h o he pollen g ains.
The i onwood ma u es and sp eads as e han he
emb yo, which p ecedes he emale game ophy e cells
by 15-20 days. Pollen, spilled om pollen g ains, si s
igh ly a he mou h o he pis il and a his ime
begins o g ow.
The emale gene a i e sphe e is o med simul aneously
wi h he male one, and his happens e y slowly (Fig. 3a-
3o). Du ing meiosis in Decembe , each o he young
lowe s e eals a pollen sac and wo co yledons. I should
be no ed ha h oughou he win e hey g ow slowly,
he lowe pa s uni e and wo nes o a ies a e o med.
The uppe ee pa o each emb yonic o ule ecomes a
column and is 3-4 imes longe han he en i e lowe
pa o he o med o a y. In mid-Ma ch, du ing he
ipening o pollen g ains, wo s yles a e o med a he
mou h o he pis il o mos o he genesis. The seeds o
he emb yo in a ma u e s a e ha e he shape o a
c assula and ha e wo di ec ions. Du ing lowe ing, he
seeds o he emb yo a e a he a chespo ial cell s age.
Bu emale game es a e eady o ecei e elemen s
capable o ep oduc ion, which begin o o m a e 2-3
weeks. The emale's well-de eloped mou h holds he ipe
pollen g ain un il he emb yo sac is ipe o ecei e i .
Wi h weak g ow h o he pollen ube on he su ace o
he pis il, he only a chespo ial cell inside he s yle is
loca ed 10-12 imes lowe han he nu cellus cell, om
which he megaspo e is o med, which unde goes
meiosis. As a esul , a e se e al mi o ic di isions, a
se en-celled, eigh -nuclea ed emale game ophy e o he
Polugonum ype is o med. The pola co e emains eady
o a long ime, only me ging wi h each o he nea
b eeding. Syne gid cells ha e a s alk-like shape, he
numbe o an ipodes is 3. The ca i y o he emb yo sac is
no e y la ge.
15
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In . J. Biosci.
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4
Fig.
3
.
Fo ma ion o he emale ep oduc i e s uc u e
a- young ui lea es (longi udinal sec ion, Decembe ), b- de elopmen o he o ule (c oss sec ion, Ma ch), c-
young o a y, q
- o a y be o e lowe ing, c - longi udinal sec ion o he o a y (Ap il), d - ma u e o a y, j -
ma u e
seed, h, i- mo he cell o mechaspo e (end o Ma ch), j- mechaspo e in e ad, l- emb yo sac
emi io (beginning o
Ap il), m
- nuclea appa a us, n- double e iliza ion (beginning o May), o- p oemb yo (4-
5 weeks a e
e iliza ion, May).
Each o a y p oduces wo ana opes, sessile seed
shoo s and in egumen . The pe iod be ween
pollina ion and ep oduc ion akes 15-20 days. The
di ision o he zygo e begins a e he endospe m has
8-12 nuclei. In pa allel wi h his, he p ocess o cell
o ma ion occu s. The i s wo-week-old emb yo
de elops acco ding o he Soland ype, he emb yo
becomes ilamen ous Kaul (1969). I should be no ed
ha emb yo di e en ia ion las s 3-4 mon hs. A he
end o Augus , he ca i y o he ge mina ing seed is
illed wi h an emb yo and consis s o a hizome, a
hypoco yl, wo s ong seeds and a shoo . The i on ee
emb yo lacks chlo ophyll. Acco ding o Masie a
(1972), plan ing esh seeds in he all gi es 80%
ge mina ion in he sp ing. I has been es ablished ha
in he i on ee, when he seeds begin o inc ease in
size, hey do no con ain ei he endospe m o emb yo.
The e o e, he emb yo sac, which is no p one o
ep oduc ion, unde goes degene a ion and pe sis s
o ou weeks a e pollina ion.
In such emb yo sacs, an ipodal cells a e always well
de eloped and esemble an egg. A he s age o
o ma ion o young ui s in İ on ee, when he
peduncles all o , he dis ance be ween he nodes
inc eases, he head o he peduncles inc eases 2-3
imes and is e ac ed in o a so , e ec s em. End ess
(1993) con i ms in his s udies such a ans o ma ion
o he cock oach du ing on ogenesis. As can be seen,
in he hi d pe iod he lowe s a e collec ed on a so
s em in a o able en i onmen al condi ions o
lowe ing plan s o he Hamamelidaceae amily.
Howe e , as a esul o he in luence o cold clima ic
condi ions, he dis ance be ween he nodes is educed
and peonies a e o med acco ding o he capi a e
ype. The i onwood ui consis s o wo seed pods.
Speaking abou he his o y o i on ees, back in he
40s o he 20 h cen u y, G ossheim (1940) classi ied
his plan as a elic wi h a wide ecological ampli ude
and Sa a o (1977) con i ms his idea in his wo k. A
p esen , his plan g ows in Lanka an, Gabala,
16
Gasimo a and Mu ado
In . J. Biosci.
202
4
Zaga ala, Abshe on, e c. I is also success ully g own in
egions o Aze baijan wi h di e en ecological condi ions
(Mamedo e al., 2016). I should be no ed ha i on ee
has a no mal g owing season, bu i s lowe ing and seed
p oduc ion a e i egula . The gene a i e o gans o İ on
ee a e cha ac e ized by low elas ici y.
Conclusion
The s udy o shoo de elopmen , lowe ing biology
and emb yology shows ha no de ia ions we e
allowed du ing hese p ocesses. The s udy o a
numbe o unique ea u es and s uc u al elemen s in
he de elopmen o İ on ee is one o he e idence
p o ing he elic na u e o his plan .
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Rohwe JG, Bi ich V. (eds) Flowe ing Plan s
.
Dico yledons. The Families and Gene a o Vascula
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G ossheim AA. 1940. Relics o Eas e n
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species le el da ed angiospe m phylogeny o
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Kaul U. 1969. Endospe m in Pa o iopsis
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Mu ado PZ, Gasimo a GCh, Namazo NR,
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Paushe a ZP. 1974. Wo kshop on plan cy ology.
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emb yos: How did we end up he e? Plan Rep od
34(4), 365-371.
Sa a o IS. 1972. I on ee Pa o ia pe sica S.A.M.
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