SEM studies on the leaf indumentum of six Melastomataceae species from Brazilian Cerrado / Microscopia eletrônica de varredura do indumento foliar de seis espécies de Melastomataceae do cerrado

2011·4.2 MB·English

by  Camilla Rozindo Dias Milanez

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Rodriguésia 62(1):203-212.2011
http://rodriguesia.jbrj.gov.br
SEM studies on the leaf indumentum
of six Melastomataceae species from Brazilian Cerrado1
MicroscopiaeletrônicadevarreduradoindumentofoliardeseisespéciesdeMelastomataceaedocerrado
Camilla RozindoDiasMilanez2&Silvia RodriguesMachado3
Abstract
The widediversity ofthcirtrichomes, which vary from simple unicellularto very complex structures, isa
remarkable characteristic in Melastomataceae. This papcr characterizes the leafindumentum ofMiconia
albicans (Sw.) Triana, M. chamissois Naudin, M.fallax DC., M. ligiistroides (DC.) Naudin, Microlepis
oleaefolia(DC.)TrianaandRhynchantheradichotomaDC.,typicalspeciesfromBraziliancerrado.Samples
collectedfromthemedianthirdofyoungandmatureleafbladeswcreprocessedfollowingtheusualscanning
electron microscopy techniques (SEM). We observed ten morphological types oftrichomes and four of
emergences.Withfivedifferenttypes,fourofwhicharereportedforthefirsttime,Rhynchantheradichotoma
isthespeciesthatpresentsthemostdiverseindumenta.Amixedtypeoftrichomeformedbyaglandularand
abranchcdnon-glandularportioncalled“lateral-gland”wasobservedinM. ligustroides.Suchnon-glandular
portionpresentsdifferentdegreesofdcvelopment.Acorrelationissuggestedbetweenthestageofdcvelopment
ofthenon-glandularportion andtheexposition tolightofthese"lateral gland”on youngleaves.
Keywords: emergence, leaf,trichome,morphology.
Resumo
Agrandediversidadedetricomas,variandodesdeestruturasunicelularesatricomasmuitocomplexoséuma
característicamarcanteemMelastomataceae.EstetrabalhocaracterizaoindumentofoliardeMiconiaalbicans
(Sw.)Triana,M.chamissoisNaudin,M.fallaxDC.,M.ligustroides(DC.)Naudin,Microlepisoleaefolia(DC.)
TrianaeRhynchantheradichotoma DC.,espéciestípicasdocerradobrasileiro. Amostrasretiradasdoterço
medianodo limbode folhasjovens e adultas foram processadas segundo técnicasusuais para microscopia
eletrônicadevarredura(MEV).Verificaram-sedeztiposmorfológicosdetricomasequatrodeemergências.
Rhynchanthera dichotoma foi aespécie que apresentou maiordiversidade de indumento, cinco, sendo que
destes, quatro foram registrados pela primeira vez. Um tipo misto de tricoma constituído por uma porção
glandulareoutranão-glandularramificada,denominadode“lateral-gland”,foiobservadoemM.ligustroides.
Aporção não-glandularapresentoudiferentesgrausdedesenvolvimento. Sugere-seumacorrelaçãoentreo
grau dedesenvolvimentodaporçãonão-glandulare aexposição àluzdestestricomasem folhasjovens.
Palavras-chave: emergência, folha,tricoma,morfologia.
W
Introduction Accordingto urdack(1986),Melastomataceae
Melastomataceae is the largest family of present the most diverse indumentum types
Myrtales,with 66gencraandapproximately4,570 among the angiosperms, even though only about
1
species, predominantly distributed in tropical and 1 1%oftheneotropical specieshavebeenstudied.
subtropical regions (Clausing & Renner2001). In The type of indumentum and lhe morphology
Brazil, itisoneofthe mostrepresentativefamilies of trichomes are important to classify
of the Cerrado flora, with about 237 species, Melastomataceae and circumscribe their genera
predominantlyshrubs (Mendonçaetal. 1998). andspecies(Wurdack 1986;Guimarães&Martins
1Partofthefirstauthorisdoctoraldissertation.ProgramadePós-graduaçãoemCiênciasBiológicas,UniversidadeEstadualPaulista.
3UniversidadeFederaldoEspíritoSanto,Depto.CiênciasBiológicas,SetordcBotânica,Av.FernandoFerrari514,29075-910,Vitória,ES,Brasil.Correspondingauthor:
[email protected]
JUniversidadeEstadualPaulista,InstitutodeBiocièncias,Depto.Botânica,C.P.510,18618-000,Botucatu,SP,Brasil.
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1997; Guimarães etal. 1999), and often allow to Material and Methods
recognize species even in a vegetative State Plant material
&
(Guimarães Martins 1997).
Studied speciesare: Miconiaalbicans (Sw.)
Studiesonleafindumentumusuallyanalyze
Triana,MiconiachamissoisNaudin,Miconiafallax
dried malure leaves of herbarium specimens,
DC.,Miconialigustroides(DC.)Naudin,Microlepis
which may lead to erroneous interpretations
oleaefolia (DC.) Triana and Rhynchanthera
sincesometrichomesarepersistemwhileothers
dichotomaDC.Theywereselectedbecauseoftheir
fali during leaf development (Uphof 1962). In
importance index at a well-preserved fragment of
addition, the distinction between the types of
cerradolocatedatthe“PalmeiradaSerraReserve”
trichomes and the structure of the indumentum
is not often clearly drawn in the literature. Such in the township of Pratânia, State of São Paulo
(22°48’50.2”Sand48°44’35.8”W),Brazil.Voucher
distinction is important in an ecological eontext
specimensweredepositedatthe Herbariumofthe
becauseitis Iikelythatenvironmenthasagreater DepartmentofBotany(BOTU),BotucatuCampus,
influenceon indumentum modificationsthan on
UniversidadeEstadualPaulista,Brazil.Eachspecies
changes in the type oftrichome (Johnson 1975).
Developmentalpatternsarenoteasilycategorized dataarelistedinTable 1
sincetrichomesareproducedon almostall plant
Study site
organs at various stages of their development.
Trichome production may continue through Thestudy site isaseasonal savannareferred
ontogeny, and as a consequence, different asCerrado(Oliveira&Marquis2002).Thisbiome
kinds oftrichomesmaybeproduced atdifferent is extremely variable in physiognomy and ranges
degreesofdevelopment(Hammond& Mahlberg from open grasslands to forests with a
1973; Johnson 1975). Evidence from the discontinuous layer of deciduous and ever-green
ontogenetic and developmental studies indicate trees with a low and shrubby growth form. Most
that pubescence is related to genetic control trees and shrubs have thick bark, twisted trunks
and environmental factors, especially to andscleromorphicleaves(Franco2002). Soilsare
availabilityofwaterregimesandradiation, since deep, strongly acid dystrophic latosols, with high
it is stimulated by red and inhibited by far-red aluminumcontents.ClimateisCwb(mesothermal
radiation, as a typical example of the with dry winters) according to Kõppen’s (1931)
phytochrome reaction (Johnson 1975; Gitz & classification. The warmest month presents an
Liu-Gitz 2003). Usually, leaves developing in average temperature inferior to 22"C, and July is
environments with high radiation tend to show the coldest and driest month ofthe year. Annual
hightrichomedensity(Crawley 1997; Sandquist rainfallis 1,534mm,withadistinctdryseasonfrom
& Ehleringer 1997). May to September (81-89 mm, respectively).
In this paper, we characterized the leaf Average(diurnal)relativehumidityisaround80%
indumentum of young and mature leaves of duringtherainyseason,butitdropsto55%during
MelastomataceaespeciesoccurringintheBrazilian the dry season when daily minimum relative
Cerrado as a framework for taxonomic and humidityreachesvaluesaround 15%.Meanannual
ecophysiological studies. temperatureisabout20,3"C.
Table 1 - Melastomataceae species studied and their respective environment at cerrado vegetation, habit and
herbariumrecord.
Species Environment Habit BOTUHerbarium
Miconiaalbicans(Sw.)Triana Cerrado stricto sensu Shrub 24,283
Miconia chamissois Naudin Borderofgalleryforest Shrub 24,284
MiconiafallaxDC. Cerradostrictosensu Shrub 24,285
Miconia ligustroides (DC.) Naudin Borderofgallery forest Shrub 24,286
Microlepisoleaefolia (DC.)Triana Borderofgalleryforest Shrub 24,287
Rhynchanthera dichotoma DC. Wet open site Sub-shrub 24,288
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Scanning electron microscopy study intercostal region ofthe abaxial surface (Fig. 2a).
(SEM) On mature leaves (Fig. 2b), they occur on both
Leaf blade samples were taken from the surfacesand aredistributed along the veins and in
medianthirdofyoungandmaturefunctionalleaves, the intercostal regions.
situated between the 2"d and 6'h nodes from the Type V: Lateral glands (Fig. 2a-e). A mixed
apical bud downwards. Materials were collected typeofhairformedbyglandularandnon-glandular
fromthreespecimensofeachspeciesandprocessed portions; lhe non-glandular portion is usually
for analysis by scanning electron microscopy branched and may present different degrees of
(SEM).Sampleswerefixedinglutaraldehyde(2.5% development (Fig. 2c-e). Always prostrate, the
with 0.1 M phosphatebuffer, atpH 7.3; ovemight glandularportion consists ofa short or long stalk
at 4°C), dehydrated in a graded alcohol series, andanelongated,biseriateglandularhead(Fig.2c-
critical-point dried (Robards 1978), mounted on e). In theintercostal regionsofthe abaxial surface
aluminumstubs,gold-coatedandexaminedwitha ofyoungleaves, hairspresentapoorlydeveloped,
Phillips515scanningelectronmicroscope. oftenbifurcatednon-glandularportion(Fig. 2c-d).
The terminology employed to describe Trichomes whose non-glandularportion is highly
indumenta follows Payne (1978), Theobald et al. developed and branched occur predominantly on
(1979),Wurdack(1986)andMentink&Baas(1992). the veins of the abaxial surface of young leaves
(Fig. 2a, e) and on the adaxial surface of mature
Resulte leaves(Fig.2b).
The analysis of the leaf surfaces of lhe TypeVI: Non-glandularemergences with a
species understudy revealedthe occurrence of 1 multiseriate wide base and piercing ápices (Fig.
differentmorphologicaltypesofindumenta: 2b,f).Theyoccuralongthemarginsofyoungand
matureleaves.
Miconiaalbicans (Sw.)Triana Fig. 1 a-b
TypeI:Non-glandular,vermiform,feiruginous Microlepisoleaefolia (DC.)Triana Fig.3a-c
trichomes(Fig. 1a-b).Theyoccuronthesuifaceof TypeVII: Dendritic,long-stalked,multiseriate
youngandmatureleaves.Onyoungleaves,theyfonn non-glandularhairs,uniformlybranched(Fig.3a).They
adense indumentum on both surfaces. On mature occur on the abaxial surface ofyoung and mature
leaves, they form a dense cover on the abaxial leavesandaredistributedoverthewholesurface.
surface (Fig. la) but are restricted to the midrib Type VIII: Dendritic, predominantly short-
regionontheadaxialsurface(Fig. 1b). stalked, non-glandular emergences, uniformly
branched(Fig.3b).Theyoccurontheadaxialsurface
Miconia chamissois Naudin Fig. 1 c-d of young and mature leaves, where they form a
Type II: Sessile to short-stalked non- dense cover.
glandularstellatetrichomes,coveredwithalveolar TypeIX: Glandular,short-stalkedhairswith
cuticle(Fig. 1c-d);theyarepresentonbothsurfaces amulticellularglandularhead(Fig.3c).Theyoccur
of young leaves, and distributed along the veins on the abaxial and adaxial surface of both young
and the intercostal regions. Mature leaves are andmature leaves.
glabrous.
Rhynchantheradichotoma(Desr.)DC.
Miconiafallax DC. Fig. 1 e-f Fig. 3d-h;4a-g
TypeIII:Sessiletoshort-stalkednon-glandular TypeX: Glandular,long-stalkedemergences
stellate trichomes (Fig. le-f); occurring on both with a pedestal base and a multiseriate, ovate
surfacesofyoungleavesandontheabaxialsurface multicellularglandularhead(Figs.3d-g;4a,0-They
ofmatureleaves, formingadensecover.Theyare are present on both surfaces ofyoung and mature
predominantlydistributedintheintercostalregions. leaves, distributed along the veins and in the
intercostal regions. Visible to the naked eye, they
Miconialigustroides (DC.)Naudin Fig.2a-f stand out by their reddish color due to abundant
TypeIV:Glandulartrichomes,predominantly anthocyanins.
short-stalked,prostrate,withanelongated,slightly TypeXI:Glandular,long-stalkedemergences
roughened,multicellularglandularhead(Fig.2a-c). with a pedestal base and a multiseriate, branched
On young leaves, they are abundant in the multicellular glandular head (Fig. 4a-d). The
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Figure 1 -a-f. LeafindumentumofMelastomataceaefromcerrado, a-b. MatureleafofMiconiaalbicans-a. abaxial
surface showing vermiform trichomes; b. adaxial surface showing trichomes restricted to midrib. c-d. Young leafof
Miconiachamissois-c.abaxialsurfaceshowingnon-glandular,stellatetrichomescoveredwithalveolarcuticle;d.detail
oftrichomes.e-f.MatureleafofMiconiafallax-e.abaxialsurfaceshowingdenseindumentum;f.detailofnon-glandular,
stellatetrichomes.Scalebars: a=66.7gm;b=64.5gm;c=71.4gm;d=48.8gm;e=91.2gm;f=62.5gm.
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Leafindumentum of Melastomataceae species hom Brazilian Cerrado 207
Figure 2 - a-f. Leafindumentum ofMiconia ligustroides- a. abaxial surface ofa young leafshowing glandular
trichomesandlateralglands;b.adaxialsurfaceofamatureleafshowingglandulartrichomes,lateralglandsandnon-
glandularemergences; c. detail ofa glandulartrichome and lateral gland; d. lateral glands showing non-glandular
portion slightly developed; e. lateral gland with a highly developed, branched non-glandular portion. The arrow
indicatestheglandularportionofalateralgland;f.detailofanon-glandularemergence.(GT=glandulartrichome;
LG=lateralgland;NE=non-glandularemergence).Scalebars:a= 194.1 pm;b=285.7pm;c=23.3 pm;d=26.7pm;
e=16.7pm;f=74pm.
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Milanez, C.R.D.&Machado,S.R.
208
Figure3-a-h.LeafindumentumofMelastomataceaefromcerrado,a-c.MatureleafoíMicrolepisoleaefolia-a.abaxial
surface showing dense indumentum composed ofdendritic, long-stalked trichomes; b. adaxial surface covered with
dendritic,non-glandularemergences;c. detailofaglandulartrichomeontheadaxialsurface. d-h. Abaxial surfaceofa
mature leafofRhynchanthera dichotoma - d. general aspect; e. detail ofthe previous figure, showing long-stalked
glandularemergences,andaglandulartrichome;f.aspectsofglandularemergence,glandulartrichome,andsessilegland
ontheintercostalregion;g.aspectsofglandularemergencewithapedestalbase,andglandulartrichomeonasecondary
vein;h.detailofanon-glandulartrichomeandaglandulartrichome.(GE=glandularemergence,GT—glandulartrichome,
NT=non-glandulartrichome; SG=sessilegland).Scalebars:a=105.3pm;b=51.3 pm,c—27.8pm,d—645.2pm,
e=166.7pm;f=47.6pm;g=91 pm;h=25.6pm.
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Leafindumentum ofMelastomataceae species from Brazilian Cerrado 209
Figure4-a-g.IndumentumabaxialsurfaceofamatureleafofRhynchantheradichotoma-a.generalaspcctshowing
avarietyoftrichomesandemergences;b.detailofalong-stalkedemergencewithabranchedglandularhcad(arrow);
c.detailofthebaseofaglandularemergence,showingstomata(arrows).Noteglandulartrichomc;d.detailofthebranched
glandularhead ofan emergence; e. glandular trichomes distributed along a secondary vein; f. glandulartrichome,
sessileglandandglandularemergencedistributedalongtheintercostalregion;g.detailofanon-glandulartrichomeanda
glandulartrichome.(GE=glandularemergence;GT=glandulartrichome;NT=non-glandulartrichome;SG=sessile
gland). Scalebars: a=400gm;b= 166.7 gm;c=38.5 gm;d= 18.9gm;e=87 gm;f=91.5gm;g=35.1 gm.
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presenceofstomataatthebaseoftheseemergences sessile glands; and short, multiseriate non-
iscommonlyobserved(Fig.4c)alongthemargins glandular trichomes. Wurdack (1986) recorded
ofyoung and mature leaves. Visible to the naked lhe occurrenceoflong-stalkedglands with thin-
eye, their reddish color is due to abundant walled headson the hypanthium and the adaxial
anthocyanins. surface of leaves of R. dichotoma. The sessile
TypeXII:Claviform,short-stalkedglandular and spherical glands observed on the surface of
hairswilhamulticellularglandularhead(Figs.3e-h; R. dichotoma leaves are similartothetrichomes
4a, c, e-g), occurring on both surfaces of young described by Metcalfe & Chalk (1950) as
andmature leaves. “bladder-like glands”. These glands are similar
TypeXIII: Spherical,sessileglands(Figs.3f; tothe pearl-likeonesobservedinPiperregnellii,
4f) present on both surfaces ofyoung and mature which were related to mucilage and protein
leaves, except on the veins. secretion (Silva&Machado 1999).
TypeXIV: Short,multiseriate,ercet(Fig.3h) A mixed type of trichome formed by a
ordeclined(Fig.4g)non-glandularhairsobserved glandular part and a branched, non-glandular
intheintercostalregionsofbothsurfacesofyoung portion similar to a lateral-gland, reported by
and matureleaves. Wurdack (1986) on the leaves of Miconia
glyptophylla and M. plumifera var. bangii, was
Discussion alsoobservedonthesurfaceofyoungandmature
Trichomes were usually denser on the leaves of Miconia ligustroides. In the present
young leaves ofall the species studied here. The work, the variable degree of ramification of
functionsandadaptivevalueoftrichomesinclude the non-glandular portion was a remarkable
protection against solar radiation, reduction of feature ofthese struetures. The young leaves of
water loss, protection against predators, and M. ligustroides occupy a vertical position, thus
physical control of materiais and metabolic exposing the abaxial surface to radiation; on this
regulation(Johnson 1975).Therefore,thepresence face, veins are very prominent and exposed to
of a dense indumentum on young leaves which radiation,whiletheintercostalareasareshadowed
arefrequentlydevoidofolhertypesofprotection, and protected from light. The intercostal areas
such as waxy deposits or lignified tissues, for Showapredominanceoflateralglandswhosenon-
example, may represent an important factor of glandularportion was little developed, while the
adaptation to the Cerrado conditions of intense veinsshowedlateralglandswithahighlybranched
solar radiation and high temperatures (Coutinho non-glandular portion. Recent studies
2002;Oliveira&Marquis2002).Ontheotherhand, demonstrated a functional link between UV
the highly reflexive adaxial surfaces ofglabrous radiation and drought that acts on the induction
mature leaves, as observed in Miconia albicans, of trichome development. UV signal perception
M. cltamissois and M.fallaxcan act analogously can lead to photomorphogenic responses as
to trichomes and reflect a large part of the trichomes differentiation which may confer
incident light, thus being of great value to adaptive advantages under abiotic stress
protectagainstexcessive insolation,desiccation conditions associated with high-light
and thermal variations (Hallé et al. 1978; environments, such as water stress (Gitz & Liu-
Ehleringer& Mooney 1984). Gitz2003). Inouropinion,thisisaveryinteresting
The results of this study corroborate species to study trichome morphogenesis.
previous reports on the wide diversity of The presence of vermiform hairs on the
indumenta and the predominance of varied and abaxial surface ofMiconia albicans leaves was
complex forms ofhairs among Melastomataceae alsoobservedby Wurdack (1986). Guimarãeset
(Solereder 1908;Metca!fe&Chalk 1950;Wurdack al. (1999) alsorecorded this type oftrichome in
1986;Guimarãesetal. 1999). Tibouchina chamissoana and T. heteromalla,
Rhynchanthera dichotoma presented a characterizing them as a tangle of smooth and
great morphological varietyoffoliarindumenta, long hairs present on the abaxial surface of the
and four morphological types of hairs are leaves.AccordingtoBaumgratz&Ferreira(1980),
described forthe first time: glandularclaviform the upper surface of M. albicans leaves is
trichomes; long-stalked glandular emergences glabrous, while its lowersurfaceispilose. In the
with branched multicellular glandular heads; present study, we also observed vermiform
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Lealindumentum of Melastomataceae species from Brazilian Cerrado 211
trichomes distributed along lhe midrib on lhe Coutinho,M.M.2002.Obiomacerrado.In:Klein,H.L.
adaxialsurfaceofinature leaves.Thesetrichomes Eugen Warminge ocerradobrasileiro, umséculo
are abundant on bolh surfaces of young leaves depois. Editora Unesp, São Paulo. Pp. 77-92.
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As for Miconia chamissois our findings Crawley,M.J.PlantEcology.AcademicPress,New
, York. Pp. 73-131.
are congruent wilh previous descriptions
Ehleringer,J.R.&Mooney,H.A. 1984.Leafhairs:effects
classifying the mature leaves of this species as
on physiological activity and adaptive value to a
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