195
On Feb ua y 6, 1999, Benjamin Elaza i Volcani died in La Jolla,
CA, USA, om kidney ailu e a he age o 84 (Fig. 1). Volcani
(bo n Wilkansky) will be emembe ed as he pionee o he
mic obiological s udies o he Dead Sea. In a sho a icle published
in Na u e in 1936 [39], Volcani ga e he i s desc ip ion o an
indigenous mic obial communi y adap ed o he ex emely ha sh
condi ions o he Dead Sea wi h i s high sal concen a ion and he
unique ionic composi ion o i s sal s. His indings we e u he
documen ed in his Ph.D. hesis submi ed o The Heb ew Uni e si y
o Je usalem in 1940 [5] and in a e iew a icle [38].
In 1939, Volcani became a membe o he Sie Ins i u e in
Reho o , la e enamed he Weizmann Ins i u e o Science; he
headed i s labo a o y o mic obiology un il 1959, when he joined
he acul y o he Sc ipps Ins i u ion o Oceanog aphy, Uni e si y
o San Diego, La Jolla, USA, whe e he emained as p o esso o
ma ine biology un il he e i ed in 1985. In his e i emen , Volcani
esumed his in es iga ions o he Dead Sea bio a. Bo les wi h old
en ichmen cul u es, da ing om he 1930s and s o ed in his
labo a o y a Sc ipps o mo e han 50 yea s, we e opened, and
hese appea ed s ill o con ain a a ie y o iable mic oo ganisms
[1, 2, 37]. Volcani’s las publica ions on he o ms o li e in he
Dead Sea appea ed in he yea o his dea h [2, 37].
The Dead Sea p esen s ascina ing challenges o he biologis
who a emp s o unde s and he biological p ocesses in he lake.
No only does he lake con ain he highes sal concen a ion
o all na u al lakes inhabi ed by li ing o ganisms, bu he
peculia ionic composi ion o he Dead Sea wa e , wi h high
concen a ions o di alen ca ions magnesium and calcium,
is highly inhibi o y e en o hose mic oo ganisms bes adap ed
o li e in he lake. These o ganisms hus li e a o nea he uppe
limi o hei ole ance owa d high magnesium concen a ions.
Since he days o Volcani’s pionee ing esea ch on he
biology o he Dead Sea ou knowledge o he lake’s bio a has
inc eased emendously ( o e iews see e.g. [18, 21, 24, 35]).
In he p esen essay we will a emp o assess he impo ance
o Volcani’s wo k in he ligh o ou p esen unde s anding o
he biological p ocesses in he Dead Sea.
Physical and chemical p ope ies o he
Dead Sea and hei long- ime changes
In he la e 1930s, when Volcani pe o med he i s mic obiological
s udies o he Dead Sea, he salini y o he lake’s wa e was lowe
han a p esen . Volcani s a ed ha a he ime he o al sal
concen a ion a he lake su ace was 269 g/l, inc easing o 327
g/l a 50 m dep h [38]. A su ey pe o med in 1959–1960 showed
he lake o be me omic ic, wi h a “pe manen ” pycnocline a a
dep h o abou 40 m [17].
Since he beginning o his cen u y he wa e balance o he
Dead Sea has been nega i e [12]. The dec ease in wa e le el,
due in pa o clima ic changes and in ensi ied by he di e sion
o esh wa e om he Jo dan i e , has led o an inc ease in
he salini y o he uppe wa e laye s. This esul ed in he
disappea ance o he pycnocline, ollowed by an o e u n o
he wa e column a he beginning o 1979. P esen ly he sal
Aha on O en1
An onio Ven osa2
1The Ins . o Li e Sciences and The Moshe Shilo
Mine a Cen e o Ma ine Biogeochemis y,
The Heb ew Uni e si y o Je usalem, Is ael
2Dep . o Mic obiology and Pa asi ology,
Fac. o Pha macy, Uni e si y o Se illa, Spain
Co espondence o:
Aha on O en. Di ision o Mic obial and
Molecula Ecology.
The Ins i u e o Li e Sciences. The Heb ew
Uni e si y o Je usalem. 91904 Je usalem. Is ael.
Tel: +972-2-6584951. Fax: +972-2-6528008.
E-mail: [email p o ec ed]
PERSPECTIVES
INTERNATL MICROBIOL (1999) 2:195–198
© Sp inge -Ve lag Ibé ica 1999
Benjamin Elaza i Volcani
(1915–1999): Six y– h ee yea s o
s udies o he mic obiology o he
Dead Sea
Fig. 1 Benjamin Elaza i Volcani (1915–1999). Pho og aph aken by A. Ven osa
in Alican e, Spain, in Sep embe 1989
con en o he Dead Sea wa e is a ound 340 g/l a all dep hs.
The mean alues o he ionic concen a ions in 1996 we e (in
mol/l): Mg2+, 1.887; Na+, 1.594; Ca22+, 0.436; K+, 0.199;
Cl–, 6.335; B –, 0.068; and SO42–, 0.005. The lake is now
holomic ic, bu me omic ic egimes ha e occu ed om
1979–1982 and om 1992–1995 as a esul o massi e in low
o esh wa e du ing unusually ainy win e s [12].
The chemical p ope ies o he lake ha e changed beyond
a simple inc ease in he o al ionic concen a ions. A he ime
o Volcani’s ea ly s udies p ecipi a ion o CaCO3we e
occasionally obse ed, causing whi ening o he wa e [3, 38].
No such e en s ha e been documen ed in ecen yea s; howe e ,
due o he nega i e wa e balance, massi e amoun s o NaCl
now p ecipi a e om he wa e column o he lake bo om as
hali e c ys als. The weigh o hali e ha has p ecipi a ed be ween
1976 and 1992 was es ima ed o be o e abou 2.550 ×106 on
[11]. The p ecipi a ion o hali e has caused an addi ional
inc ease in he al eady ex emely high a io o di alen o
mono alen ca ions o he Dead Sea wa e .
Pho osyn he ic mic oo ganisms in he
Dead Sea
F om en ichmen cul u es se up wi h Dead Sea wa e and
sedimen samples, Volcani succeeded in g owing a a ie y o
pho osyn he ic mic oo ganisms. These include euka yo ic
lagella e algae (Dunaliella sp.), well adap ed o he high
salini ies p e ailing a he ime in he uppe wa e laye s, and
an uniden i ied g een lagella e wi h a single lagellum ha
g ew a 1–27% sal , wi h sligh g ow h a 30% [5, 6, 38].
Di e en ypes o cyanobac e ia we e also ound ha g ow a
lowe sal concen a ions (up o abou 18%) [38].
Al hough indigenous li e was disco e ed in he Dead Sea
as ea ly as 1936, he i s quan i a i e assessmen s o he mic obial
communi y densi ies in he lake we e made only as la e as
1963–1964 [14]. I is now clea ha a small Dunaliella species
(designa ed in he pas as Dunaliella i idis o Dunaliella pa a)
is he main o only p ima y p oduce in he lake. The g een
lagella e wi h a single lagellum isola ed by Volcani ha e ne e
been obse ed in he lake since, no ha e been cyanobac e ia
epo ed o occu in signi ican numbe s. Cyanobac e ia, howe e ,
can nowadays be ound in abundance in hype saline sp ings on
he lake’s sho e, such as Hamei Mazo (a sp ing wi h a salini y
o abou 169 g/l ha was subme ged a he ime o Volcani’s
s udies, when he wa e le el was highe han a p esen ) [22].
Dunaliella was eco ded o occu in su ace wa e in high
numbe s in 1964 (up o 4 ×104cells/ml). F om 1980 onwa ds a
comp ehensi e sampling and moni o ing p og am has been ca ied
ou , esul ing in an ex ensi e da a se co e ing algal densi ies o e
he seasons a di e en dep hs. Blooms o Dunaliella cells we e
obse ed du ing he pe iod 1980–1981 and again in 1992–1994:
up o 8.8 ×103cells/ml we e coun ed in 1980 [27] and 1.5 ×104
cells/ml and maybe e en highe densi ies in 1992 [23, 30]. The
appea ance o Dunaliella in he Dead Sea was igge ed by he
massi e ain loods in he win e s o 1979–1980 and 1991–1992
ha caused a signi ican dilu ion o he uppe wa e laye s, and
ini ia ed a me omic ic s a e o he lake. Dunaliella cells we e
ound only in he mixed epilimnion abo e he pycnocline. In mos
samples examined, he dis ibu ion o he algae was uni o m om
he wa e su ace down o he pycnocline. Howe e , du ing he
pe iod Augus –Oc obe 1992 cells we e ound concen a ed as a
“deep chlo ophyll maximum” a a dep h o 7–10 m nea he
pycnocline [30].
Labo a o y simula ions ha e shown ha a dilu ion o he uppe
wa e laye s wi h mo e han 10% esh wa e is equi ed o ini ia e
a mass de elopmen o he alga, p o ided ha all essen ial nu ien s
a e a ailable. Inpu o su icien phospha e is also necessa y, as
phospha e is he limi ing nu ien in he Dead Sea.
When he salini y becomes oo high o he Dunaliella cells,
hey apidly disappea om he wa e column. Du ing he
decline o he dense algal bloom in he summe o 1992,
Dunaliella cells we e obse ed o o m cys s ha sank o he
bo om o he lake [30].
Halophilic A chaea
Among he mic oo ganisms isola ed by Volcani ha a e bes
adap ed o he ex emely high sal concen a ions o he Dead Sea
a e a numbe o ed halophilic A chaea belonging o he amily
Halobac e iaceae. Haloa cula ma ismo ui, i s desc ibed by
Volcani as Fla obac e ium (o Halobac e ium) ma is-mo ui,
was epo ed o g ow a NaCl concen a ions om 18% up o
sa u a ion [5, 10, 38]. The gene ic name Halobac e ium was i s
used by Volcani in his Ph.D. hesis, and he consolida ed he
nomencla u e o he genus in his con ibu ion o he 7 h edi ion
o Be gey’s Manual o De e mina i e Bac e iology in 1957 [10].
The o iginal cul u e o Halobac e ium ma ismo ui has no been
p ese ed, bu a simila s ain now desc ibed as Haloa cula
ma ismo ui was isola ed om he lake in he la e 1960s [13],
and a o mal desc ip ion o he species was based on his isola e
[29]. In addi ion, Volcani also isola ed a “ ed mic ococcus”,
p obably being a Halococcus s ain [5, 38].
Addi ional no el halophilic A chaea ha e been isola ed
om he Dead Sea by o he in es iga o s; hese include
Halo e ax olcanii, named a e Volcani [16], Halo ub um
sodomense [20], and Halobaculum gomo ense [21].
Recen ly a numbe o bo les con aining en ichmen cul u es,
da ing om he 1930s and s o ed in Volcani’s labo a o y a he
Sc ipps Ins i u ion o Oceanog aphy in La Jolla o mo e han
50 yea s, we e opened, and he mic oo ganisms hey con ained
we e analyzed. Molecula analysis showed ha hese cul u es
con ained a a ie y o a chaeal species, some o hem no el [1,
37]. This sugges ha ou unde s anding o he biodi e si y o
he lake is s ill a om comple e.
Quan i a i e assessmen s o he a chaeal communi y in he
Dead Sea showed ha halophilic A chaea a e o en p esen in
196 INTERNATL MICROBIOL Vol. 2, 1999 O en, Ven osa
e y high concen a ions: be ween 2.3 ×106and 8.9 ×106cells/ml
we e coun ed in 1963–1964 [14]; up o 1.9 ×107cells/ml we e
ound in he lake’s su ace laye s in he summe o 1980 [19],
and in he sp ing o 1992 a maximum communi y densi y o
3.5 ×107cells/ml was eached [26]. The dense communi ies
o he A chaea ich in ca o enoid pigmen s impa ed a eddish
colo o he Dead Sea wa e bo h in 1980 and in 1992 [19, 23,
26], and a eddish hue had been also obse ed in 1964 [14].
De elopmen o a chaeal blooms is co ela ed wi h he occu ence
o la ge numbe s o Dunaliella, he p ima y p oduce in he lake,
which supplies he necessa y o ganic ma e ial. Glyce ol, p oduced
and accumula ed by Dunaliella as an osmo ic s abilize , is
p obably one o he main sou ces o ca bon ha suppo he
de elopmen o he a chaeal communi y.
Li le is known on he con ibu ion o he di e en gene a and
species o halophilic A chaea o he communi y in he Dead Sea.
Du ing he 1992 bloom, pola lipid analyses we e pe o med o
ob ain in o ma ion on he ypes o A chaea p esen . The lake’s
biomass con ained one majo glycolipid ( he sul a ed diglycosyl
die he lipid S-DGD-1), whe eas he die he de i a i e o
phospha idylglyce osul a e (PGS) was absen [25]. These
obse a ions may sugges ha he dominan A chaeon in he Dead
Sea a he ime belonged o he genus Halo ub um o o
Halobaculum. No signi ican amoun s o hose speci ic glycolipids
we e ound ha would ha e indica ed a massi e p esence o
Haloa cula ma ismo ui in he biomass a he ime.
Halophilic and halo ole an ae obic
Bac e ia
In addi ion o halophilic A chaea, Volcani isola ed a numbe o
halophilic o halo ole an Bac e ia om he Dead Sea. Two o
hese ha e been p ese ed and desc ibed as new species:
Ch omohalobac e ma ismo ui [36] (o iginally desc ibed as
Ch omobac e ium ma is-mo ui [5, 38]), and Halomonas
halmophila [4] (o iginally Fla obac e ium halmephilum [5, 38]).
Ano he in e es ing isola e, which un o una ely was no p ese ed,
is Pseudomonas hales o gus [5, 38]. F om he old en ichmen
cul u es, a new halophilic Bacillus species, Bacillus ma ismo ui,
was ecen ly isola ed [2]. Non-halophilic ep esen a i es o he
genus Bacillus we e also ound in he Dead Sea [5, 38]. These
we e e med “halo esis an ” by Volcani. These o ganisms p obably
su i ed exposu e o he hype saline b ines o he Dead Sea in
he o m o esis an endospo es, and canno be conside ed o
belong o he indigenous mic obio a o he lake. To hono Volcani’s
achie emen s in he ield o halophilic Bac e ia, he bac e ial
genus Volcaniella was named a e him [33].
Anae obic mic oo ganisms in he
sedimen s
F om he bo om sedimen s o he Dead Sea, Volcani isola ed
a numbe o halophilic anae obic bac e ia [7, 38]. These include
e men a i e bac e ia— hey e men glucose o lac ose and
g ow a 25% sal —and deni i ie s. Se e al ae obic halophilic
A chaea, including Haloa cula ma ismo ui, can g ow by
deni i ica ion. Un o una ely, he e men a i e isola es ob ained
by Volcani we e no p ese ed. Only in he ea ly 1980s we e
new isola es o halophilic e men a i e Bac e ia ob ained om
he Dead Sea and om o he hype saline lakes, and he g oup
(o de Haloanae obiales) appea ed o be an ex emely in e es ing
one [34]. Dead Sea isola es belonging o his o de a e
Halobac e oides halobius [32], O enia ma ismo ui, and
Spo ohalobac e lo e ii [34]. Spo ohalobac e lo e ii was
named a e he F ench mic obiologis M. L. Lo e , who in
he las decade o he 19 h cen u y isola ed a numbe o
pa hogenic non-halophilic ep esen a i es o he genus
Clos idium om Dead Sea mud [15]. These p obably had
su i ed as esis an endospo es, and should hus be designa ed
halo esis an bac e ia acco ding o Volcani’s classi ica ion.
P o ozoa
In en ichmen cul u es incuba ed o e y long pe iods (mon hs
o yea s), Volcani ound wo ypes o p o ozoa: a cilia e and
a dimas igamoeba [8, 9, 38]. La e s udies ha e ne e
asce ained he p esence o p o ozoa in he Dead Sea ecosys em
o hei possible impo ance in egula ing communi y densi ies
o mic oo ganisms in he lake. Vi uses, howe e , may be
implica ed in he decline o a chaeal and algal blooms: many
i us-like pa icles we e obse ed upon examina ion o wa e
samples in he elec on mic oscope [28].
Cilia e and amoeboid p o ozoa we e abundan in he
hype saline (169 g/l o al sal ) sul u sp ing o Hamei Mazo ,
which, a he ime o Volcani’s s udies, was subme ged below
he lake su ace, and now is exposed on he sho e [22]. Thus,
he p o ozoa cul u ed by Volcani migh ha e de i ed om less
saline ecosys ems in he Dead Sea a ea.
Epilogue
The apidly changing Dead Sea can be conside ed an
in e es ing la ge-scale expe imen in mic obial e olu ion and
adap a ion wi h espec o highe and highe di alen ca ion
concen a ions. A he ime o w i ing, a chaeal communi y
densi ies in he lake we e e y low, and no Dunaliella cells
we e obse ed a all. The A chaea and he es ing s ages o
he algae in he bo om sedimen s we e wai ing o hei
oppo uni y o mul iply when condi ions would become
sui able again. Li e in he Dead Sea in i s p esen s a e hus
depends p ima ily on hose a e e en s o abundan ain all
in he ca chmen a ea ha lead o he o ma ion o an
epilimnion su icien ly dilu ed. The e e inc easing salini y
o he Dead Sea wa e and he concomi an ly inc easing
197
B. E. Volcani (1915–1999) INTERNATL MICROBIOL Vol. 2, 1999
ela i e concen a ions o inhibi o y di alen ca ions, along
wi h he inc easing ex en in which excess ainwa e in he
ca chmen a ea is di e ed o ag icul u al use, sugges ha
such mic obial bloom e en s may become a e and a e .
Volcani’s s udies in he la e 1930s p o ide us wi h aluable
in o ma ion on he ypes o mic oo ganisms ound in he Dead
Sea du ing a pe iod in which he o e all salini y was much lowe
han oday, and he uppe wa e laye had less han 80% o i s
p esen -day sal concen a ion. Un o una ely, no quan i a i e
in o ma ion was collec ed by Volcani, and almos 30 yea s had
o pass un il he i s enume a ions o mic oo ganisms in he lake
we e pe o med [14]. Ne e heless, he in o ma ion o be ound
in Ben Volcani’s legacy, consis ing bo h o his publica ions and
o hose bo les o en ichmen cul u es p ese ed om he 1930s,
is o he u mos impo ance o unde s and he mic obial ecology
o he Dead Sea, no only in he pas bu also in he p esen .
Re e ences
1. A ahal DR, Dewhi s FE, Pas e BJ, Volcani BE, Ven osa A (1996)
Phylogene ic analysis o some ex emely halophilic A chaea isola ed
om Dead Sea wa e , de e mined on he basis o hei 16S RNA
sequences. Appl En i on Mic obiol 62:3779–3786
2. A ahal DR, Má quez MC, Volcani BE, Schlei e KH, Ven osa A (1999)
Bacillus ma ismo ui sp. no ., a new mode a ely halophilic species om
he Dead Sea. In J Sys Bac e iol 49:521–530
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desc ip ion o Halomonas halmophila (NCMB 1971T). In J Sys Bac e iol
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