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STRUCTURAL DIVERSITY AND ANATOMICAL
ADAPTATIONS OF LEAVES IN BRASSICACEAE SPECIES
G.S. Tu sinbaye a
Associa e p o esso , The B anch o ASTU in Tashken egion, Uzbekis an
h ps://doi.o g/10.5281/zenodo.17674830
Abs ac . This s udy p esen s a compa a i e ana omical desc ip ion o lea s uc u e in
se e al species o he amily B assicaceae Bu ne , including Alyssum dasyca pum, A.
u kes anicum, A. szo i sianum, Dip ychoca pus s ic us, Goldbachia lae iga a, Hymenolobus
p ocumbens, Isa is minima, and I. iolascens. Mic oscopic analysis e ealed dis inc in e speci ic
a ia ions in ichome ype, s oma al dis ibu ion, and mesophyll o ganiza ion. Mos s udied
species possess amphis oma ic lea es wi h isola e al-palisade mesophyll and scle enchyma ized
ascula bundles, e lec ing xe omo phic and heliophilous adap a ions. Di e ences we e
obse ed in palisade hickness, deg ee o pubescence, and epide mal cell mo phology. These
ana omical cha ac e is ics p o ide aluable diagnos ic ea u es o axonomy and ecological
adap a ion s udies wi hin he B assicaceae amily.
Keywo ds: b assicaceae; lea ana omy; isola e al-palisade mesophyll; amphis oma ic
lea es; xe omo phic adap a ion; Alyssum; Isa is; Hymenolobus p ocumbens; Dip ychoca pus
s ic us; Goldbachia lae iga a; epide mal s uc u e; ichomes; s oma al ypes; ascula bundles.
In oduc ion
The B assicaceae Bu ne (C uci e ae) amily includes nume ous species adap ed o
di e se en i onmen al condi ions, pa icula ly a id and semi-a id egions. Acco ding o C.R.
Me cal e and L. Chalk (1957), he lea es o B assicaceae species a e p edominan ly do si en al
bu may also be isola e al-palisade. The ichomes a e s uc u ally di e se, and s oma a a e
ypically anisocy ic on bo h sides o he lea . Idioblas s con aining my osin, which exhibi ed o
pu ple colo a ion upon coagula ion, se e as a diagnos ic ea u e o he amily. E.I. Volko a
(1960) in es iga ed he ana omy o en epheme al B assicaceae species om Kope dagh and he
Moscow egion, no ing an inc ease in mesomo phy unde in oduc ion condi ions. La e , N. Rao
and I. Inamda (1983) examined lea mo phology and ena ion in 35 B assicaceae species,
desc ibing hem as simple, al e na e, and p edominan ly c aspedod omous. G.F. Begbae a (1995)
cha ac e ized he mesophyll s uc u e o Isa is species, emphasizing hei ana omical and
nu i ional alue (Ka imo e al., 1965). Howe e , de ailed in o ma ion on he ana omical s uc u e
o dese B assicaceae species emains limi ed. The e o e, his s udy aims o p o ide a compa a i e
ana omical analysis o selec ed B assicaceae species o iden i y diagnos ic and adap i e ea u es
ela ed o hei ecological condi ions.
Aim o he esea ch: The s udy aims o in es iga e he ana omical ea u es o lea es in
selec ed B assicaceae species and o e eal diagnos ic and adap i e s uc u al ai s ela ed o hei
ecological dis ibu ion, pa icula ly in a id and semi-a id habi a s.
Objec o he esea ch: The objec s o his s udy a e he lea es o eigh B assicaceae
species—Alyssum dasyca pum, A. u kes anicum, A. szo i sianum, Dip ychoca pus s ic us,
Goldbachia lae iga a, Hymenolobus p ocumbens, Isa is minima, and Isa is iolascens—analyzed
h ough compa a i e ana omical me hods.
2. Ma e ials and Me hods
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The s udy examined lea es o Alyssum dasyca pum, A. u kes anicum, A. szo i sianum,
Dip ychoca pus s ic us, Goldbachia lae iga a, Hymenolobus p ocumbens, Isa is minima, and I.
iolascens.
F esh lea samples we e collec ed om na u al popula ions and ixed in a 70% e hanol
solu ion. Tempo a y mic oscopic slides we e p epa ed using s anda d pa a in embedding and
mic o ome sec ioning echniques. Pa ade mal and ans e se sec ions we e s ained wi h sa anin
and as g een o isualize cell walls and issues. Obse a ions we e made unde a ligh mic oscope.
Mo phological ea u es—including epide mal s uc u e, ichome ype, s oma al dis ibu ion,
mesophyll o ganiza ion, and ascula bundle a angemen —we e analyzed and compa ed among
species.
3. Resul s and Discussion
3.1. Alyssum dasyca pum. Lea es a e simple, b oad-lanceola e (6–8 mm × 4–6 mm),
en i e, and pubescen wi h 5–8- ayed s ella e hai s. The epide mis is single-laye ed; cells ha e
sligh ly sinuous walls wi h hickened ou e laye s. The lea is amphis oma ic, wi h o al,
hemipa acy ic, anisocy ic, and a ely anomocy ic s oma a. The mesophyll is isola e al-palisade
and loose; palisade issue is single-laye ed on bo h sides, while he spongy pa enchyma has h ee
laye s. Vascula bundles (25–30) a e scle enchyma ized.
3.2. Alyssum u kes anicum Lea es a e simple, oblong (9–11 mm × 2–3 mm), densely
pubescen wi h 16- ayed s ella e hai s. The epide mis is single-laye ed, wi h wa y cell walls and
sligh ly hickened ou e walls. Lea es a e amphis oma ic; s oma a a e elonga ed, anisocy ic, and
e acy ic. The mesophyll is isola e al-palisade wi h 2–3 laye s o palisade cells on he adaxial side
and one laye on he abaxial side. The spongy issue has 4–6 laye s. The ascula bundles (19–21)
a e scle enchyma ized.
3.3. Alyssum szo i sianum Lea es a e oblong-poin ed (7–9 mm × 4–6 mm), pubescen
wi h 8–16- ayed s ella e hai s. The epide mis is single-laye ed, wi h amphis oma ic su aces.
S oma a a e anomocy ic and hemipa acy ic. The mesophyll is isola e al-palisade, wi h 2–3 laye s
o palisade cells on he adaxial side, 1–2 laye s on he abaxial side, and 3–4 laye s o spongy issue
in be ween. The cen al bundle is weakly scle enchyma ized.
3.4. Dip ychoca pus s ic us Lea es a e simple, oblong-linea wi h sha p se a ions,
co e ed wi h simple and glandula hai s. The epide mis is single-laye ed wi h wa y (adaxial) and
sinuous (abaxial) walls. The lea is mainly do si en al wi h h ee laye s o palisade cells and 5–6
laye s o spongy issue. Vascula bundles (20–22) a e s ongly scle enchyma ized.
3.5. Goldbachia lae iga a Lea es a e hea -shaped a he base, pe iola e (8–10 mm × 1.5–
2 mm), and spa sely se a ed. They a e co e ed wi h simple unicellula hai s. The lea is
amphis oma ic wi h small anomocy ic and hemipa acy ic s oma a. The mesophyll is isola e al-
palisade wi h 2–3 laye s o palisade cells on bo h sides and 6–7 laye s o spongy pa enchyma.
Vascula bundles a e scle enchyma ized, su ounded by a pa enchyma ous shea h.
3.6. Hymenolobus p ocumbens Lea es show p onounced he e ophylly: basal lea es a e
ly a ely lobed, while uppe lea es a e en i e and sessile. Epide mal cells ha e wa y walls, and
s oma a a e anisocy ic and hemipa acy ic. The mesophyll is isola e al-palisade (277.9 µm hick)
wi h 3 palisade laye s adaxially, 2 abaxially, and 6 spongy laye s be ween. Vascula bundles a e
scle i ied, indica ing xe omo phic adap a ion.
3.7. Isa is minima Lea es a e simple, wi h inely se a ed ma gins. The epide mis is single-
laye ed, amphis oma ic, and consis s o wa y-walled cells. S oma a a e anomocy ic and
anisocy ic. The mesophyll is isola e al-palisade, wi h 2–3 palisade laye s adaxially and 1 abaxially,
and 6–7 loose spongy laye s. All ascula bundles a e scle enchyma ized.
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3.8. Isa is iolascens Lea es a e b oad a he base, ape ing owa d he apex, and bea hin
ilamen ous and hook-shaped ichomes. An hocyanin pigmen a ion occu s in abaxial epide mal
cells. The lea is amphis oma ic, wi h nume ous anomocy ic and anisocy ic s oma a. The
mesophyll is do si en al wi h 2 palisade and 7–8 spongy laye s. Vascula bundles (24–26) a e
hea ily scle i ied.
All s udied B assicaceae species sha e se e al diagnos ic ea u es: amphis oma ic lea es,
non-sunken s oma a, s ella e o simple ichomes, and single-laye ed epide mis wi h hickened
ou e walls. Mos species exhibi isola e al-palisade mesophyll, e lec ing xe omo phic and
heliophilous adap a ions o a id condi ions. The genus Alyssum demons a es no able in e speci ic
a ia ion: A. dasyca pum has b oad-lanceola e lea es wi h mode a e pubescence, A. u kes anicum
displays elonga ed lea es and a high palisade a io (63%), and A. szo i sianum has hicke
mesophyll wi h ewe scle enchyma ized bundles.
Species such as Hymenolobus p ocumbens and Isa is minima show ansi ional ea u es
be ween mesomo phic and xe omo phic ypes, sugges ing adap i e lexibili y. The p esence o
an hocyanin in I. iolascens may enhance p o ec ion agains high sola adia ion. O e all, he
ana omical di e si y obse ed among hese species indica es e olu iona y esponses o habi a
condi ions, con i ming ha lea s uc u al ai s in B assicaceae a e aluable diagnos ic and
ecological indica o s.
Family B assicaceae Bu ne In he comp ehensi e wo k by C.R. Me cal e and L. Chalk
(1957), i is no ed ha he lea es o species in he B assicaceae amily a e p edominan ly
do si en al bu can also be isola e al-palisade. The ichomes a e highly di e se. The s oma a a e
anisocy ic on bo h sides o he lea . Idioblas s con aining my osin a e p esen in all o gans, which
u ns ed o pu ple upon coagula ion, making i a use ul diagnos ic ea u e. E.I. Volko a (1960)
s udied he s uc u e o 10 epheme al species, including B assicaceae species, in he condi ions o
Kope dagh and he Moscow egion. She no ed he di e si y o lea ypes and an inc ease in
mesomo phy unde in oduc ion condi ions in he Moscow egion.
N. Rao and I. Inamda (1983) desc ibed he mo phology and ena ion o he lea es o 35
B assicaceae species, including species o he genus Alyssum. The lea es a e simple and al e na e,
bu he shape o he blade is highly a iable. The ena ion is p ima ily c aspedod omous. G.F.
Begbae a (1995) desc ibed he ype o mesophyll in he lea es o wo Isa is species. Isa is species
a e aluable o hei nu i ional con en , con aining all essen ial amino acids and signi ican
amoun s o ca bohyd a es (Ka imo e al., 1965). Thus, in o ma ion abou he lea s uc u e o
dese species in he B assicaceae amily is sca ce.
Alyssum dasyca pum: The lea is simple, b oad-lanceola e, en i e, and smoo hly ansi ions
in o a pe iole. The lea blade is 6-8 mm long and 4-6 mm wide, e enly pubescen on bo h sides
wi h 5-8- ayed s ella e hai s. The epide mis is single-laye ed, wi h 4-5-sided cells wi h sligh ly
sinuous walls in pa ade mal sec ions. Thei ou e walls a e sligh ly hickened. On he abaxial side,
he epide mal cells a e smalle and lowe han on he adaxial side in c oss-sec ion. The lea is
amphis oma ic. The s oma a a e supe icial, o al, hemipa acy ic, anisocy ic, and a ely
anomocy ic. The mesophyll is isola e al-palisade and loose. The palisade issue is single-laye ed
on bo h sides, and he spongy issue is h ee-laye ed. The ascula bundles a e scle enchyma ized,
wi h 25-30 bundles in c oss-sec ion.
Alyssum u kes anicum: The lea is simple, oblong, and almos sessile. The blade is 9-11
mm long and 2-3 mm wide, en i e, and densely pubescen wi h 16- ayed s ella e hai s, dense on
he adaxial side. The epide mis is single-laye ed. In pa ade mal sec ions, he cells a e 4-5-sided
wi h sligh ly sinuous walls on he adaxial side and wa y walls on he abaxial side. The ou e walls
a e sligh ly hickened. On he adaxial side, he epide mal cells a e la ge and alle han on he
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abaxial side. The lea es a e amphis oma ic. The s oma a a e elonga ed-o al, anisocy ic, and
e acy ic. The mesophyll is isola e al-palisade, wi h 2-3 ows o palisade cells on he adaxial side
and one ow on he abaxial side. The palisade issue occupies mos o he mesophyll olume, wi h
he heigh o he palisade cells being wice hei wid h. The spongy issue is 4-6 laye s hick. The
ascula bundles a e nume ous, wi h 19-21 in c oss-sec ion, and a e scle enchyma ized.
Alyssum szo i sianum: The lea is simple, oblong-poin ed, wi h a blade 7-9 mm long and
4-6 mm wide, en i e, and pubescen on bo h sides wi h 8-16- ayed s ella e hai s (mainly 8- ayed
on he uppe side and 16- ayed on he lowe side). The epide mis is single-laye ed. The lea es a e
amphis oma ic. In pa ade mal sec ions, he walls o he adaxial epide mis a e sligh ly wa y, while
he abaxial walls a e sinuous. The s oma a a e anomocy ic and hemipa acy ic, la ge bu ewe in
numbe on he adaxial side o he lea . The mesophyll is isola e al-palisade, wi h 2-3 ows o
palisade cells on he adaxial side and 1-2 ows on he abaxial side. Be ween hem a e 3-4 ows o
loose spongy cells. The in e cellula spaces a e la ge h oughou he mesophyll. The ascula
bundles a e small, he la e al bundles a e no scle enchyma ized, and he cen al bundle has a ew
scle enchyma cells on he abaxial side and is su ounded by pa enchyma.
Fig. 1. S uc u e o he lea o Alyssum dasyca pum:
a – ex e nal appea ance;
b – c oss-sec ion diag am;
c – dend oid ichomes;
d – de ail o he mesophyll;
e – adaxial epide mis;
– abaxial.
B
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Fig. 2. S uc u e o he Alyssum u kes anicum lea :
a – ex e nal appea ance;
b – c oss-sec ion diag am;
c – ichomes;
d – de ail o he mesophyll;
e – adaxial epide mis;
– abaxial.
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Fig. 3. S uc u e o he lea o Alyssum szo i sianum:
a – ex e nal appea ance;
b – c oss-sec ion diag am;
c – de ail o he mesophyll;
d– main ein;
e – dend oid ichome;
– adaxial epide mis;
g – abaxial.
The lea es o Alyssum szo i sianum, A. u kes anicum, and A. dasyca pum sha e he
ollowing common ai s: hey a e co e ed wi h s ella e hai s, amphis oma ic (ha ing s oma a on
bo h sides), he s oma a a e non-imme sed, and he adaxial epide mal cells a e la ge han he
abaxial ones. The mesophyll is o he isola e al-palisade ype. Howe e , he lea es o species in
he Alyssum genus di e in he ollowing diagnos ic ea u es:
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Alyssum szo i sianum is cha ac e ized by a lanceola e lea shape and hick mesophyll.
Alyssum u kes anicum has an elonga ed lea shape, a hinne lea , and mo e p onounced
palisade issue (palisade a io – 63%).
A. dasyca pum has a b oad-lanceola e lea shape.
In he on ogeny o plan s, xe omo phic and heliomo phic s uc u es o assimila ing o gans
in ensi y: om do si en al co yledons o isola e al-palisade lea es.
Dip ychoca pus s ic us: The lea is simple, oblong-linea , wi h sha p se a ions, and
co e ed wi h simple and glandula hai s on a 4-celled s alk. The epide mis is single-laye ed. The
su ace cells a e la ened, wi h wa y cell walls on he adaxial side and sinuous on he abaxial side.
The s oma a a e nume ous, especially on he abaxial side, anomocy ic, hemipa acy ic, anisocy ic,
and non-imme sed.
The mesophyll is do si en al, wi h h ee ows o wide palisade cells on he adaxial side
and loose spongy cells on he abaxial side. The spongy issue consis s o 5-6 laye s. The ascula
bundles (main and 20-22 la e al) a e scle enchyma ized.
Goldbachia lae iga a: The lea is lamina , hea -shaped a he base, pe iola e, spa sely
se a ed along he edges, 8-10 mm long, 1.5-2 mm wide, ape ing a he ip. I is co e ed wi h
simple unicellula hai s on a wo-celled base. The epide mal cells in pa ade mal sec ions a e
la ened and sligh ly wa y; in c oss-sec ions, hey a e swollen wi h a hickened ou e wall. The
lea is amphis oma ic. The s oma a a e small, nume ous, anomocy ic and hemipa acy ic, and non-
imme sed. The mesophyll is isola e al-palisade, wi h 2-(3) ows o palisade cells on bo h sides and
6-7 ows o spongy cells in he cen e . The median ascula bundle is la ge han he la e al ones
and is su ounded by a scle enchyma ous shea h.
Hymenolobus p ocumbens: The lea es exhibi homoblas ic he e ophylly. The basal lea es
a e al e na e, numbe ing 3-5, up o 1.5 cm long and 0.8 cm wide, ly a ely pinna e-lobed wi h 5-8
lobes. The lowe s em lea es a e 1.5-1.8 cm long and 0.5 cm wide, dissec ed wi h 4-5 lobes. The
middle lea es a e 1.2-1.5 cm long and 0.4-0.5 cm wide, less dissec ed, oblong, wi h an elonga ed,
sligh ly poin ed apex and a d awn-ou base, sho -pe iola e o sessile. The uppe lea es a e 0.5-1
cm long and 0.1-0.3 cm wide, en i e, and sessile. In c oss-sec ions o he middle lea es, he
epide mis is single-laye ed, and he ou e walls a e hickened (4.7 μm). Table 1
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Fig. 4. S uc u e o he lea o Dip ychoca pus s ic us:
a – ex e nal appea ance;
b – c oss-sec ion diag am;
c – ichome;
d – de ail o he mesophyll;
e – main ein; – adaxial epide mis; g – abaxial.
Fig. 5. S uc u e o he Goldbachia lae iga a lea :
a – ex e nal appea ance;
b – c oss-sec ion diag am;
c – de ail o he mesophyll;
d – adaxial epide mis;
e – abaxial;
– main ein;
g – ichomes.
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On he pa ade mal sec ion, he epide mal cells a e elonga ed, wi h wa y walls on bo h sides
o he lea . The s oma a a e anisocy ic and hemipa acy ic, less equen ly anomocy ic, non-sunken,
and andomly o ien ed. The mesophyll is isola e al-palisade, wi h a hickness o 277.9 µm. On he
adaxial side, he e a e 3 laye s o palisade cells, and on he abaxial side – 2 laye s wi h la ge
in e cellula spaces and 6 laye s o spongy cells in be ween. The palisade coe icien is 45%, and
he palisade index o he cells is 2. The median ascula bundle has 10-15 essels, wi h 10-12
small la e al bundles. All bundles a e scle i ied due o he hickening o he walls o he phloem
pa enchyma (Table 1; Fig. 3 ). In he on ogeny o H. p ocumbens, he e is a change in he
adap a ion o assimila ing o gans owa ds an inc ease in heliophilous and xe omo phic ai s,
exp essed in he ansi ion om do si en al mesophyll in co yledons o isola e al-palisade
mesophyll in lea es. Howe e , he s uc u e o ege a i e o gans also has mesomo phic ea u es:
wa y epide mal cells, la ge in e cellula spaces in he lea mesophyll, and nume ous small non-
sunken s oma a.
Isa is minima. The lea is simple, wi h inely se a ed edges and a p ominen ein on he
abaxial side. The epide mis is single-laye ed. The epide mal cells a e la ened, hei walls a e
wa y, highe on he adaxial side, wa y and la ge on he abaxial side. The lea is amphis oma ic.
The s oma a a e nume ous, small, anomocy ic, a ely hemipa acy ic and anisocy ic, and non-
sunken. The mesophyll is isola e al-palisade, wi h 2-(3) ows o palisade cells on he adaxial side
and 1 ow on he abaxial side. The spongy issue is 6-7-laye ed, loose, wi h la ge in e cellula
spaces. The median ascula bundle p o udes om bo h he adaxial and, e en mo e so, he abaxial
sides. The e a e 10 la e al ascula bundles on he ans e se sec ion. All bundles a e scle i ied
(Fig. 4).
Conclusion
The compa a i e ana omical analysis o eigh B assicaceae species—Alyssum dasyca pum,
A. u kes anicum, A. szo i sianum, Dip ychoca pus s ic us, Goldbachia lae iga a, Hymenolobus
p ocumbens, Isa is minima, and I. iolascens—demons a es ha lea s uc u al ai s wi hin his
amily se e as eliable indica o s o ecological adap a ion and axonomic di e en ia ion. Despi e
species-speci ic di e si y, all s udied axa sha e a se o diagnos ic ea u es cha ac e is ic o
B assicaceae: amphis oma ic lea es, non-sunken s oma a, a single-laye ed epide mis wi h sligh ly
hickened ou e walls, and he p esence o simple o s ella e ichomes. These a ibu es
collec i ely e lec adap a ion o en i onmen s wi h high sola adia ion and luc ua ing mois u e
a ailabili y.
Mos species exhibi isola e al-palisade mesophyll, a xe omo phic and heliophilous ai
ha enhances pho osyn he ic e iciency unde a id and semi-a id condi ions. Va ia ions in palisade
hickness, ichome densi y, and deg ee o ascula bundle scle enchyma iza ion u he e eal
e olu iona y adjus men s o mic ohabi a condi ions. The genus Alyssum demons a es
p onounced in e speci ic di e en ia ion, pa icula ly in lea shape, palisade a io, and deg ee o
pubescence, con i ming i s high ecological plas ici y. Species such as Hymenolobus p ocumbens
and Isa is minima display ansi ional ea u es be ween mesomo phic and xe omo phic ypes,
indica ing lexible esponses o mois u e a ailabili y ypical o dese and oo hill ecosys ems. The
p esence o an hocyanin pigmen a ion in I. iolascens sugges s an addi ional pho op o ec i e
mechanism agains excessi e i adiance.
O e all, he s uc u al di e si y o lea es ac oss he examined species e lec s complex
e olu iona y esponses o a id habi a s and suppo s he ecological signi icance o ana omical ai s
in B assicaceae. These indings no only e ine diagnos ic c i e ia o species iden i ica ion bu
also con ibu e o unde s anding adap i e s a egies in dese and semi-dese lo as, whe e
ana omical modi ica ions play a key ole in su i al unde en i onmen al s ess.
