JULY, 1958] HELMINTHOLOGICA L SOCIET Y 13 3
On th e Morpholog y of th e Coffe e Root-Kno t Nematode,
Meloidogyne exigua Goeldi, 1887*
LUIZ GONZAG A E. LORDELL O AND ADIE L PAES IjEM E ZAMIT H
Escola Superio r de Agricultur a "Luiz de Queiroz, " Universit y of S . Paulo ,
Piracicaba , Brazil
The firs t recor d on nematode s attacking root s of coffe e trees in Brazi l was
publishe d by Jober t in 1879 . Som e years later , Goeld i (1887) stated that the
primar y cause of a diseas e affectin g coffee plantation s at th e so-calle d Prov -
inc e
of Ri o d e Janeir o was th e nematod e referre d to b y Jobert . At tha t time ,
Goeld i erecte d the ne w genu s Meloidogyne, and describe d Meloidogyne exigua
as th e typ e species . Sinc e then , author s hav e mad ' reference s to root-kno t
nematode s found infectin g coffee trees in severa l Sout h America n countries ,
and in th e U . S . A. , at th e Ne w Yor k Botanica l Garde n (Chitwood , 1949;
Lordello , 1953; Taylor , Dropki n & Martin, 1955; etc.) . In 1949 , Chitwoo d
publishe d
his well-know n paper on th e root-kno t nematodes , proposin g the
invalidatio n of th e genu s Meloidogyne, whic h had bee n synonymize d with
Heterodera Schmidt , 187 1 by previou s authors .
Recently ,
the writer s examine d root s of coffe e trees grow n in th e Ribeira o
Preto area , in th e Stat e of S . Paulo , Brazil , and foun d M. exigua attackin g
all th e decayin g plants . Experiment s for recuperatin g old coffe e orchard s
have bee n conducte d at on e o f th e plantation s infecte d by M . exigua, bu t th e
tree s did no t respon d as expected . Actually , preliminar y survey s sugges t tha t
M. exigua i s a n importan t detrimen t to coffe e productio n in th e Ribeira o
Preto region . M. exigua produce s somewha t smal l gall s on th e root s of coffe e
trees. Thes e gall s may b e easil y overlooked , particularl y if th e materia l col -
lecte d is no t protecte d agains t desiccation . Necrotic areas are als o to b e see n
on th e roots . The tree s foun d infecte d belon g to th e following " varieties of
Coffea arabica L. : red Bourbon , yello w Bourbo n and "Mund o Novo."
M. exigua is a littl e know n root-kno t nematod e species. The descriptio n
by Goeld i is adequat e to plac e the genus , but i s erroneou s in man y respects .
In addition , Chitwoo d coul d not giv e an entirel y satisfactor y redeseriptio n be-
caus e the materia l he ha d fo r stud y was i n poo r condition . For thos e rea -
sons ,
the observation s carrie d out wit h the abundan t materia l collecte d at
Ribeira o Preto are her e presente d as a contributio n to th e knowledg e of
M. exigua, th e coffe e root-kno t nematode .
Meloidogyne exigua Goeldi, 188 7
EGGS : The egg s are ellipsoidal , 73.4-88.7 micron s lon g and 38.3-44. 4 mi-
cron s w 7ide (fig \, J). Observatio n mad e on egg s containin g larva e did no t
offe r any indicatio n that the firs t larva l moul t take s plac e insid e the egg .
PREPARASITI C LARVAE : Th e wor m shape d bod y taper s to bot h extremities , more sharpl y posteriorly , endin g in a n elongate d tail . Hea d bearin g one pos t
labia l
annule ; stylet bulb s weakl y develope d and rathe r compresse d longi -
tudinally . Cuticle annulated , the annulatio n being muc h less eviden t tha n in
adults . Middl e bul b of oesophagu s ovoid. Oesophagea l gland s rathe r long'
(fig. 1, H , an d I) . Th e nerv e ring encircle s the oesophagea l isthmu s jus t be-
low th e middl e bulb. Intestina l cell s filled with coagulate d substances , forming
*The author s wis h to expres s their best thank s to Dr . Herman o V. d e Arruda , of th e Estaca o Experimenta l de Ribeira o Preto, S. Paulo , Brazil , who supplie d the m wit h severa l samples of root s of coffe e trees disfigure d by M . e.riyuu.
Copyright © 2011, The Helminthological Society of Washingto n
134 PROCEEDING S OF TH E [VOL . 25, No . 2
Fig . 1.— Meloidogyn e exigua Goeldi, 1887: A. Oesophagea l regio n of th e male ; B. Hea d of th e male ; C. Posterio r end o f male ; D. Style t of male ; E. Latera l field of male ; F. Oesophagea l regio n of th e female ; G. For m of th e femal e body ; H. Oesophagea l regio n of preparasiti c larva; I. Posterio r end o f preparasiti c larva; J. Egg .
Copyright © 2011, The Helminthological Society of Washingto n
JULY, 1958] HELMINTHOLOGICA L SOCIETY
mor e
or les s roun d and characteristi c bodies . Anal openin g sometime s hard
to locate . Phasmid s not seen . Latera l fields 3. 1 micron s wide , mad e up o f fou r
incisures .
Measurement s (in microns) . Length: 333.5-358.0 ; width: 13.7-15.3 ; tail:
44.4-46.0 ; stylet: 9.2; middl e bul b of oesophagus : 10.7 x 7.7 ; lengt h of oesophagus : 78.0-82.6; anal diameter : 7.7-9.2; a = 22.2-26.0 ; b = 4.2-4.4 ;
c = 7.3-7.8.
FEMAL E : Th e whitis h female s hav e well define d neck and wer e obtaine d
from th e smooth-walle d cavitie s of th e roo t tissue s (fig . 1, G) . A numbe r of
dea d female s w7as als o obtaine d durin g dissectio n of th e galls . Thes e female s
wer e violaceou s
in colo r and ha d rathe r resistan t cuticle , lookin g lik e wal l of
cyst s of th e Heterodera species .
Head pointed , style t straigh t or slightl y curved , provide d wit h smal l
knobs . Cana l
of th e oesophagu s strongl y walle d fro m the beginnin g of th e
orga n unti l it enter s the valve s of th e middl e bulb , wher e it become s rathe r
weak .
A constrictio n of th e surroundin g tissue s is see n at th e poin t of unio n
of th e oesophagea l cana l wit h the middl e bulb , the latte r being ovoi d to
spherica l and quit e stron g (fig . 1, F) . Latera l fields an d excretor y por e not
Tig. 2.— Meloidogyn c exigua Goelcli, 1887: Perinea ! patterns of adul t females .
Copyright © 2011, The Helminthological Society of Washingto n
136 PROCEEDING S OF TH E [VOL . 25, No . 2
seen . Ovarie s convoluted , usuall y filled with oocytes . The perinea l pattern
agreed with that figure d b y Chitwoo d (1949 ) and Taylor , Dropki n & Martin
(1955) for th e species . The arch is lo w an d slightl y flattene d an d th e latera l
line s poorl y
define d and bordere d by folde d and broke n striae (fig. 2).
Measurement s (in microns) . Length: 387.5-496.0; width: 279.0-372.0 ;
stylet: 10.7; distanc e from the orific e of th e dorsa l glan d to bas e of stylet :
4.6-7.7 ; oesophagu s middl e bulb: 30.6-33. 6 x 24.5-26.0 .
MALE : Body worm-lik e in shape , taperin g more abruptl y toward anterio r
end. Th e head , mor e frequentl y obtaine d in latera l or sublatera l view, is
slightl y set of f fro m neck . Labia l annule wide and rathe r flat i n latera l view;
post labia l annul e withou t any striatio n in dorso-ventral , as wel l as i n latera l
view. Cuticl e strongl y annulated . Lateral fields beginnin g a little back of th e
leve l of th e style t knob s and extending * to tai l end . The y are usuall y mad e
up o f fou r incisures , but i n severa l male s additiona l incisures were counted .
The transvers e stria e of cuticl e may cu t th e oute r band s of th e fields , a s
illustrate d (fig . 1, E) . At abou t the middl e of th e body , the field s ar e 7.6-9. 2
micron s wide .
Meloidogyne male s usuall y hav e a twiste d body . M. exigua males , how -
ever ,
are no t twisted , thu s constitutin g an exceptio n amon g the root-kno t
nematodes already studie d by th e writers . Actually , othe r specie s previousl y
investigate d hav e male s
to whic h the twis t of th e bod y was calculate d to b e 9 0
degree s (Lordello , 1956, and 1956a) . Twistin g may be considere d as bein g
adaptativ e
in character , since it i s suppose d that the male s have to rol l thei r
body aroun d the femal e bod y whe n copulatio n take s place .
Stylet strong , the knob s bein g ovoi d and ver y pronounced . The nerv e ring
encircle s the isthmu s jus t belo w the middl e bul b (fig . 1, A) . Intestina l cells
filled with coagulate d bodie s of differen t forms. Excretor y por e wel l defined ,
located at 96.0-130. 0 micron s fro m the hea d end . All th e male s studie d hav e
two wel l develope d testes , whic h may b e reflexe d or completel y outstretched ,
usuall y reachin g
the oesophagea l region . Spicule s arcuate d ventrally ; guber-
naculu m well defined . Phasmid s very smal l (fig . 1, C) .
In th e populatio n handled , two type s of male s were found , whic h differe d
in th e distanc e betwee n the openin g of th e dorsa l oesophaga l glan d and th e
style t knobs . In on e type , tha t distanc e is aroun d 3.0 microns , whil e in th e
othe r the orific e is to o clos e to th e knob s to measure . Sinc e no difference s
were observe d in th e othe r stage s of th e parasite , the tw o mal e type s wer e
considere d as individua l variation s withi n the sam e specie s (fig . 1, B , an d D) .
In latera l and sublatera l views , the ampulla of th e amphid s may b e mor e
or les s easil y located , clue to it s circula r outline . The writer s prefe r to us e
the ter m ampulla fo r suc h amphidia l structure in th e genu s Meloidogyne, in - stea d of th e wor d "cheek " introduce d by Chitwoo d (1949 ) which , accordin g
to th e writers ' opinion , is quit e inadequate . As alread y show n by Alle n
(1952) , Cobb (1924 ) did not use the wor d "cheek " to designat e the organ
mentione d above, but state d tha t "th e amphid s are protecte d by cheeks. " As Allen (1952 ) does , the writer s understan d tha t Cob b referre d to th e wel l de- velope d lateral lip s of th e Meloidogyne species.
Measurement s (in microns) . Length: 832.3-1,092.4 ; width: 26.0-46.0 ;
stylet: 18.4-19.9 ; distance fro m the openin g of th e dorsa l glan d to bas e of stylet : 0.0-3.0 ; style t knobs : 4.0-6.1 x 3.0; middl e bul b of oesophagus : 15.3
x 9.2 ; tail: 6.1-10.0 ; spicules (measure d on chor d of thei r arc) : 20.0-26.0 ;
gubernaculum : 7.7; heigh t of head : 3.1-4.6 ; a = 23.8-32.0 ; b = 8.1-8.9 ; c = 95.8-110.0 .
Copyright © 2011, The Helminthological Society of Washingto n
JULY, 1958] HELMINTHOLOGICA L SOCIETY 13 7
LITERATURE CITED
ALLEN , M. W . 1952 . Observation s on th e genu s Meloidogyne Goelcl i 1887. Proc. Helminth . Soc. Wash . 19: 44-51 . CHITWOOD , B. G . 1949 . "Root-kno t nematodes"—par t I. A revisio n of th e genu s Meloidogyne Goelcli , 1887. Proc . Helminth . Soe . Wash . 16: 90-104 . COBB, N. A . 1924 . The amphid s of Caconema (nom . iiov.) and o f othe r nemas . Jour. Parasitol . 11: 118-120 . GOELDI , B. A . 1887 . Belatori o sobre a molesti a do cafeeir o na. Proviiici a do Bi o d e Janeiro . Arch. Mus. Nac. Eio d e Janeir o 8: 7-123 , year: 1892 . JOBERT , C. 1878 . Sur un e maladi e du eafeie r observe e au Bresil . C. E . Acad . Sci. Pari s 87: 941-943 . LORDKLLO , L. G . E . 1953 . Contribuica o ao conheciment o dos nematodeo s do sol o de alguma s regioes do Estad o de S . Paulo . Thesis, Escola Superio r de Agricultur a "Luiz de Queiroz, " Universit y of S . Paulo , 75 pp. , 5 pis. , Piracicaba , Brazil. . 1956. Meloidogyne inornata sp. n., a seriou s pest of soybea n in th e Stat e of S. Paulo , Brazi l (Nematoda, Heteroderidac). Rev. Brasil , Biol. 16: 65-70 . . 1956a . Nematoidc s que parasita m a soja na regia o de Bauru . Bra- gantia 15: 55-64 . TAYLOR , A. L. , V. II . DROPKIN , and G . C . MARTIN . 1955. Perinea ! patterns of root - knot nematodes . Phytopatholog y 45: 26-34 .
Alkalin e Phosphatas e in the Trematod e Excretory System *
WILLIA M H. COI L
The functio n of th e excretor y system amon g the platyhelmint h worms is ye t
to b e understood ; however, both Beave r (1929 ) and Wille y (1934 ) hav e pub-
lished note s on th e morpholog y of th e flam e cells in trematodes . There are goo d
reason s
to believ e that this syste m function s in bot h osmoregulatio n and ex -
cretion . Kromhou t (1943) showe d that , in th e cas e of th e turbellarians , the
excretor y syste m is mos t highl y develope d in th e fres h wate r form s and least
in th e marin e species . On th e othe r hand , Buchana n (1931) found that tur-
bellarian s gai n weigh t by imbibitio n in distille d wate r and los e weigh t in
sal t water , thus showin g that this syste m can be overworke d unde r extrem e
conditions .
In contras t to this , man y trematode s can pas s fro m fres h wate r
to solution s wit h variou s osmoti c pressure s during the variou s stages of thei r
life cycles . The motio n of th e flam e cells can b e stimulate d by salin e (Beave r
1929 )
or urine . Thi s coul d be eithe r a respons e to th e ion s of differen t in-
organi c salts or a respons e to a n osmoti c change . The presenc e of concretion s
in th e excretor y bladder s of som e trematode s woul d lea d one t o believ e thes e
are th e resul t of excretor y action .
Severa l year s
ago whe n I wa s attemptin g to wor k out th e comple x excre -
tor y syste m
of som e gorgoderi d trematodes , it occurre d to m e tha t ther e mus t
be a n easie r way tha n the stud y of livin g material . Thi s led t o th e specu -
latio n that alkalin e phosphatase , present in th e excretor y system s of man y
othe r animals , migh t be presen t in th e excretor y syste m of trematodes . At
that tim e this postulat e was tested , but fo r som e reason , unknow n to me , n o
evidenc e was foun d to suppor t it. Recently , while studyin g a gorgoderi d
cercaria (close to Gercaria sphaerocerca Miller , 1936) from the gill s of a
sphaerii d bivalve , it wa s note d that the daughte r sporocyst s possessed un-
usuall y large flam e cells and associate d capillaries . It wa s though t that this
larg e materia l might wor k bette r tha n previou s specimens .
The livin g materia l was fixe d i n chille d aceton e (below 0°C) and the n
placed ,
in acetone , in a refrigerato r for 24 hours . Imbeddin g was carrie d out
*TtVo m the Departmen t of Zoology , Universit y of Nebraska , Study No. 303 , supporte d by a Grant-in-Ai d from the Universit y of Nebrask a Research Council .
Copyright © 2011, The Helminthological Society of Washingto n
