INTERNATIONAL JOURNAL O F
SYSTEMATIC
BACTERIOLOGY
Vol. 20, No. 4
October 1970
Copyright 1970, Iowa State University P r e s s
pp. 339-344
PROGRESS AND PROBLEMS IN BACTERIAL SYSTEMATICS
E r w i n F. L e s s e l
A m e r i c a n Type C u l t u r e Collection
Rockville, Maryland, U. S. A.
ABSTRACT. C o n s i d e r a b l e p r o g r e s s h a s b e e n m a d e i n b a c t e r ial systematics, particularly in characterization
a n d i n a p p r o a c h e s t o d e t e r m i n i n g t h e r e l a t i o n s h i p s of
bacteria.
Nonetheless r e a l progress is difficult bec a u s e m u c h of t h e o l d e r l i t e r a t u r e i s i n a d e q u a t e i n
t e r m s of t h e p r e s e n t s t a t e o f k n o w l e d g e a n d b e c a u s e
o r i g i n a l s t r a i n s do not e x i s t w h e r e w i t h s e n s e c a n be
m a d e o u t of t h e o l d e r l i t e r a t u r e .
It i s felt that bact e r i a l s y s t e m a t i c s could be placed on a f i r m foundation
i f the following steps w e r e taken:
1) r e v i s e t h e B a c teriological Code so that i n o r d e r for a n a m e to be
validly published a prescribed minimal description
w o u l d b e r e q u i r e d a s w o u l d d e p o s i t i o n of t h e t y p e s t r a i n
i n a c u l t u r e c o l l e c t i o n f r o m w h i c h it w o u l d b e a v a i l a b l e ;
2 ) establish a date in the future a s the starting date
f o r n a m e s of b a c t e r i a , c o n s e r v i n g , h o w e v e r ,
those
n a m e s of r e c o g n i z a b l e t a x a f o r w h i c h t h e r e a r e a d e quate descriptions and type strains (if the organism i s
c u l t i v a b l e ) , a n d 3) w h e r e i n d i c a t e d , a c q u a i n t a u t h o r s
and editors with the requirements for valid publication
and with sound taxonomic practices,
_ _ _ _ _ _ _ _ - _ - -
B a c t e r i a l s y s t e m a t i c s e m b r a c e s the c h a r a c t e r i z a t i o n , c l a s s i f i c a t i o n ,
identification and naming of b a c t e r i a . In t h e s e a r e a s a g r e a t d e a l of
p r o g r e s s has been m a d e f r o m the time b a c t e r i a w e r e first d i s c o v e r e d t o
the p r e s e n t , especially during t h e p a s t twenty-five y e a r s . However,
d e s p i t e t h e advances that have been m a d e , the c u r r e n t s t a t e of b a c t e r i a l
s y s t e m a t i c s l e a v e s m u c h t o be d e s i r e d , and the situation b e c o m e s inc r e a s i n g l y w o r s e a s new t a x a a r e d e s c r i b e d and named and as m o r e and
m o r e invalid s t a t e m e n t s a p p e a r i n t h e l i t e r a t u r e . In t h a t which follows,
s o m e mention will be made of r e c e n t advances in t h e s y s t e m a t i c s of bact e r i a , but t h e e m p h a s i s will be on t h e o b s t a c l e s to r e a l p r o g r e s s and on
t h e ways i n which t h e s e o b s t a c l e s c a n be r e m o v e d o r circumvented.
G r e a t s t r i d e s have been m a d e not only i n t h e equipment and methods
u s e d t o d e t e r m i n e the c h a r a c t e r i s t i c s which have been routinely u s e d i n
taxonomy but a l s o in the s e a r c h f o r new a t t r i b u t e s t o b e t t e r show the relationships of the b a c t e r i a . Among the l a t t e r a r e t h e p r e s e n c e o r a b s e n c e
of a n u c l e a r m e m b r a n e , t h e c y t o p l a s m i c m e m b r a n e s y s t e m , t h e n u m b e r
and a r r a n g e m e n t of flagella, t h e c h e m i c a l composition of t h e c e l l wall,
t h e appendages on s p o r e s , metabolic pathways and products, t h e DNA
b a s e r a t i o , DNA homology, and c o m p a r a t i v e c h e m i s t r y of e n z y m e s with
identical functions.
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In addition t o m o r e and improved data on the c h a r a c t e r i s t i c s of t h e
b a c t e r i a , c o m p u t e r s a r e now available to facilitate o u r handling of t h e s e
data. R e g a r d l e s s of whether o r not we want t o weight c e r t a i n c h a r a c t e r s ,
p r o g r a m s c a n be written which will enable us to c o m p a r e in a consistent
fashion l a r g e n u m b e r s of s t r a i n s and taxa. I t should be s t r e s s e d h e r e
that, as applied t o b a c t e r i a l classification, t h e computer i s a tool, not a
concept o r a philosophy. It s o happened that t h e f i r s t u s e of t h e computer in b a c t e r i a l classification was in connection with the application of
Adanson's theory, proposed in the l a t t e r half of the 18th Century, which
maintains t h a t e a c h c h a r a c t e r i s t i c i s of equal weight i n d e t e r m i n a t i o n of
r e l a t e d n e s s . Unfortunately t h e r e a r e many taxonomists who equate the
computer with Adansonian theory, and who, not subscribing to t h i s
theory, will have nothing to do with the computer. They should r e a l i z e
t h a t t h e computer c a n be a n extension of t h e i r minds and c a n include all
the bias they w i s h and that, if they a r e dealing with l a r g e n u m b e r s of
s t r a i n s o r l a r g e v o l u m e s of data, t h e function of the computer i s t o expedite t h e i r work. One of the contributions computer taxonomy h a s m a d e
is that m o s t taxonomists now r e a l i z e that in o r d e r b e t t e r to " s e e f f bact e r i a and to visualize t h e i r relationships, b a c t e r i a should be looked a t in
t e r m s of a l a r g e n u m b e r of a t t r i b u t e s r a t h e r than f r o m the n a r r o w viewpoint of a few c h a r a c t e r i s t i c s with which a taxonomist m a y be enamored;
a m u c h b e t t e r p e r s p e c t i v e of the b a c t e r i a i s obtained by looking a t t h e i r
t o t a l biological p i c t u r e r a t h e r than at a few a r b i t r a r i l y s e l e c t e d c h a r a c t e r i s t i c s j u s t a s t h e a s t r o n a u t s obtained a much better p e r s p e c t i v e of t h e
planet E a r t h , indeed of m a n himself, when they recently viewed t h e E a r t h
f r o m the proximity of the moon.
Now r e g a r d l e s s of whether o r not one supplements h i s inbuilt, biologic a l computer with a m e c h a n i c a l one to facilitate h i s computations and t o
analyze h i s data, it i s , of c o u r s e , n e c e s s a r y that the data be c o m p a r a b l e
i f valid conclusions a r e to be reached. If one u s e s his own data, t h e r e
should be no problem. If, however, one i s using information obtained
from a v a r i e t y of s o u r c e s , then valid c o m p a r i s o n s might become difficult,
i f not impossible, to make. C l e a r l y many of the older d e s c r i p t i o n s of
b a c t e r i a a r e e x t r e m e l y confusing and c r e a t e g r e a t difficulty not only bec a u s e of insufficient information when c o m p a r e d to m o d e r n d e s c r i p t i o n s
but a l s o because it is v e r y difficult t o a s s e s s the reliability of t h e data.
Valid c o m p a r i s o n of data becomes possible, and a g r e a t d e a l e a s i e r , if
detailed d e s c r i p t i o n s of o r suitable r e f e r e n c e s to the methods used to
obtain t h e data a r e given.
In addition t o being reliable, it i s e s s e n t i a l that the data on the c h a r a c t e r i s t i c s of a b a c t e r i u m be sufficient to p e r m i t the identification of the
bacterium. Enough information should be given, 1) to place the o r g a n i s m
i n its p r o p e r taxonomic niche, and 2 ) t o s e p a r a t e it f r o m closely r e l a t e d
o r g a n i s m s . Unless information of t h i s type i s provided in the d e s c r i p t i o n
of a newly n a m e d species, it m a y be i m p o s s i b l e to d e t e r m i n e t o which
o r g a n i s m the n a m e applies, and the n a m e would then be illegitimate a s a
nomen dubium. It probably would be helpful i f , in those c a s e s w h e r e
possible, the inclusion i n the d e s c r i p t i o n of each newly named b a c t e r i u m
of a m i n i m a l list of c h a r a c t e r i s t i c s significant to t h e classification of
b a c t e r i a w e r e m a d e p a r t of the r e q u i r e m e n t s for t h e valid publication of
n a m e s . Such a l i s t might include the G r a m and a c i d - f a s t reactions,
morphology, mode of reproduction, mode of motility, i f any, number and
position of flagella, i f any, relationships to oxygen, t e m p e r a t u r e and s a l t ,
and t h e type of energy-yielding metabolism. In connection with t h i s , it is
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interesting to note that nearly 6 y e a r s ago a l i s t of c h a r a c t e r i s t i c s r e commended f o r inclusion in the description of streptomycetes which
appear in patent applications was published in the International Bulletin
of Bacteriological Nomenclature and Taxonomy, 13(3):169- 170 (1963).
In the absence of adequate descriptions, cultures on which the original
descriptions were based a r e invaluable in helping taxonomists to understand the literature; these cultures a r e also very useful in updating older
descriptions. However, one disadvantage with maintaining bacteria in
s e r i a l subculture i s that a s actively growing cells they a r e subject to
mutation and selection; furthermore, the m o r e subculturing required,
the g r e a t e r the chances for contamination to occur.
Fortunately it was discovered many y e a r s ago that bacteria can be
preserved in a viable and stable condition for long periods by freeze drying; however, it was not until the 1940's that this method of preserving
bacteria gained wide acceptance. Within the past 10 y e a r s it has been
found that bacteria can be preserved m o r e effectively when frozen and
stored at the t e m p e r a t u r e of liquid nitrogen than when maintained in the
f r e e z e - d r i e d state.
Now the fact that bacteria can be preserved in a viable and stable
condition fo2 long periods has great implications in bacterial systematics.
It means that living cultures can s e r v e a s fixed standards, i. e. a s nomenclatural types, for taxonomic groups. F o r many y e a r s the?;a<teria w e r e
regarded a s plants, and their nomenclature was governed by the Botanical Code. Plants a r e classified and identified primarily on morphologic a l grounds, and dead specimens of plants s e r v e adequately a s r e p r e sentatives of nomenclatural types; living specimens, which a r e subject
to change, w e r e not permitted by the Botanical Code to s e r v e a s r e p r e sentatives of nomenclatural types. However a s the science of bacteriology developed, it soon became apparent that living cultures were indispensible to the complete characterization and to the classification and
identification of bacteria. It was primarily for the reason that the
Botanical Code did not permit living cultures to s e r v e a s nomenclatural
types that bacteriologists formulated a code of nomenclature for the bact e r i a , which was published initially in 1947. To be s u r e , the Bacteriological Code i s not perfect. However, during the past 10 y e a r s Dr. R. E.
Buchanan and his staff have put the Code through rigorous paces while
preparing the text of the Index Bergeyana and supplementary material,
and most of the problems and deficiencies a r e recognized. It now r e mains for the International Committee on Nomenclature of Bacteria to
effect a suitable revision of the Code, and towards this end an ad hoc
committee was constituted a little over 6 months ago by the Judicial Commission of the International Committee on Nomenclature of Bacteria.
In addition to revising the Bacteriological Code, t h e r e i s another step
that, if taken, should aid considerably the efforts to improve bacterial
systematics. It would appear that many of the problems in systematic
bacteriology could be avoided i f editors of biological journals were r e minded of o r made aware of the nomenclatural-type concept and the r e quirements for the valid publication and legitimacy of names of bacteria.
It is suggested, therefore, that a committee or committees of qualified
individuals be formed to a s s i s t editors of biological journals by providing
them with information and guidelines relating t o bacterial nomenclature
and perhaps by offering their s e r v i c e s on reviewing papers on systematics.
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R e g a r d l e s s of how highly we r e g a r d m o d e r n d e s c r i p t i o n s of b a c t e r i a ,
it is conceivable that in t h e future s o m e of t h e m m a y be quite inadequate
because of the l a c k of c e r t a i n bits of information. It is, t h e r e f o r e , e s s e n t i a l t h a t c u l t u r e s of type s t r a i n s be available s o that d e s c r i p t i o n s can
be updated and valid c o m p a r i s o n s made. Many, if not m o s t of the probl e m s i n b a c t e r i a l s y s t e m a t i c s would disappear i f c u l t u r e s of the type
s t r a i n s of a l l of t h e named s p e c i e s w e r e available. It s e e m s r e a s o n a b l e ,
t h e r e f o r e , to m a k e one of the r e q u i r e m e n t s for the valid publication of a
s p e c i e s o r s u b s p e c i e s n a m e t h e designation of t h e type s t r a i n and the
deposition of a c u l t u r e of the type s t r a i n in a c u l t u r e collection f r o m
which the s t r a i n would be available. F o r o r g a n i s m s which cannot be
cultivated, adequate d e s c r i p t i o n s o r i l l u s t r a t i o n s would have to suffice.
C u r r e n t l y one of t h e r e q u i r e m e n t s f o r the establishment of a neotype
s t r a i n i s that t h e s t r a i n be deposited i n a c u l t u r e collection f r o m which
it would be available; t h i s r e q u i r e m e n t should now be m a d e p a r t of t h e
conditions f o r the v a l i d publication of new s p e c i e s and s u b s p e c i e s n a m e s .
P e r h a p s the m o s t i m p o r t a n t and useful concept in b a c t e r i a l s y s t e m a t i c s is that of t h e nomenclatural type. It i s by m e a n s of the nomenclat u r a l type t h a t we a r e able t o d e t e r m i n e the application of n a m e s . T h e
n o m e n c l a t u r a l type i s that constituent e l e m e n t of a taxon to which t h e
n a m e of the taxon i s permanently attached. The nomenclatural type of a
genus is a s p e c i e s and that of s p e c i e s o r subspecies i s a s t r a i n , which
m a y be r e p r e s e n t e d by a living c u l t u r e o r by a description o r a n i l l u s t r a tion. A type s t r a i n mukt be one of t h e s t r a i n s on which the o r i g i n a l description w a s based. If none of the original s t r a i n s i s extant, t h e n a
l a t e r i s o l a t e which a g r e e s with the o r i g i n a l d e s c r i p t i o n m a y be designated
a s a neotype. According to the nomenclatural type concept, a named
s p e c i e s can be d e s c r i b e d a s consisting of the type s t r a i n together with a l l
o t h e r s t r a i n s which closely r e s e m b l e t h e type s t r a i n . If a t s o m e t i m e it
i s decided to divide the s t r a i n s which c o m p r i s e a named s p e c i e s into two
o r m o r e groups and to recognize e a c h group a s a named s p e c i e s , the
g r o u p which contains the type s t r a i n of the o r i g i n a l s p e c i e s m u s t r e t a i n
the n a m e of this s p e c i e s ; f o r e a c h of the other groups a new s p e c i e s n a m e
m u s t be given and a type s e l e c t e d f r o m the included s t r a i n s , F u r t h e r m o r e any study o r any s t a t e m e n t about a named taxon m u s t take into cons i d e r a t i o n t h e n o m e n c l a t u r a l type of that taxon. F o r example i f one
wanted to d i s c u s s the genus Corynebacterium, the type s p e c i e s , C. -dipht h e r i a e , cannot be ignored for it is the s p e c i e s to which the g e n e r i c n a m e
C o r y n e b a c t e r i u m is permanently attached, and it i s t h e only s p e c i e s
which belongs to t h e genus C o r y n e b a c t e r i u m without question,
Nomenclatural types a r e useful in the determination of synonymies.
N a m e s based on the s a m e type a r e objective synonyms; n a m e s based on
different types, which types a r e r e g a r d e d a s s i m i l a r enough t o belong to
the s a m e taxon, a r e subjective synonyms. F o r example, when a s p e c i e s
i s t r a n s f e r r e d f r o m genus to genus, the r e s u l t a n t n a m e s a r e objective
synonyms, In the c a s e of two independently named s p e c i e s , e a c h with its
own type, synonymy of the two n a m e s i s d e t e r m i n e d by c o m p a r i s o n of t h e
types; if the types a r e r e g a r d e d a s s i m i l a r enough to belong to the s a m e
s p e c i e s , the n a m e s a r e r e g a r d e d a s subjective synonyms because the r e l a t e d n e s s of the types i s a p e r s o n a l opinion.
The m a i n purpose, t h e r e f o r e , of nomenclatural types is t o provide a
fixed r e f e r e n c e point for scientific n a m e s s o that a given name will always
r e f e r t o the s a m e taxonomic group. C l e a r l y " s p e c i e s " i s a m a n - m a d e
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BACTERIOLOGY
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concept, and the l i m i t s of s p e c i e s a r e constantly in a s t a t e of flux due
not only to evolutionary p r o c e s s e s but a l s o to the many differences in
human opinion. I t i s e a s y t o s e e how the concept of a given s p e c i e s could
change o v e r the y e a r s i f t h e r e w e r e not s o m e fixed point to r e l a t e the
n a m e to. T h e r e a r e , in fact, s e v e r a l documented c a s e s w h e r e t h e s p e c i e s d e s c r i p t i o n applied to a n a m e eventually changed s o much that t h e
m o d e r n d e s c r i p t i o n does not a g r e e a t a l l with the original d e s c r i p t i o n ;
in addition, t h e r e a r e i n s t a n c e s w h e r e a n a m e h a s been used with s o
many different meanings t h a t complete confusion h a s resulted.
The p r i m a r y purpose of type s t r a i n s , then, i s to s e r v e a s fixed r e f e r e n c e s to which the n a m e s of s p e c i e s a r e permanently attached. Type
s t r a i n s have the s p e c i a l function of determining the applications of n a m e s ,
and they a r e not intended to be typical of the s p e c i e s ; i n f a c t it i s quite
difficult t o d e t e r m i n e exactly what i s a typical o r a v e r a g e s t r a i n of a
s p e c i e s . Even if a typical s t r a i n of a s p e c i e s could be d e t e r m i n e d , it
coule only r e l a t e t o a given population a t a given t i m e ; changes in the
population a r e constantly o c c u r r i n g , and we would be faced with t h e i m possible t a s k of selecting a never-ending s e r i e s of typical s t r a i n s . Suff i c e it to s a y t h a t a typical s t r a i n , if indeed it c a n be recognized a s such,
h a s a function different f r o m that of a type s t r a i n : w h e r e a s the f o r m e r is
intended a s a n example of the kind of s t r a i n m o s t frequently encountered
in a population, t h e l a t t e r s e r v e s as a permanent, fixed s t a n d a r d in the
application of n a m e s . Both type s t r a i n s and typical s t r a i n s c a n s e r v e
adequately a s points around which o t h e r s t r a i n s c a n be c l u s t e r e d . Howe v e r t h e type s t r a i n holds t h e advantage because it does not change with
time.
As stated previously, one a s p e c t of b a c t e r i a l s y s t e m a t i c s i s nomenc l a t u r e . When an author wishes t o recognize a new taxon h e m u s t give a
scientific n a m e to the taxon i n accordance with the internationally a c cepted r u l e s of nomenclature. According to the r u l e of p r i o r i t y , the
e a r l i e s t available name o r specific epithet m u s t be used. I t i s incumbent
then f o r a n author of a new name, indeed for any author who u s e s a s c i entific name, t o d e t e r m i n e whether t h e r e a r e any e a r l i e r n a m e s o r specific epithets for the taxon he i s dealing with. If done p r o p e r l y , t h i s
entails a n e n o r m o u s amount of work, for t h e r e a r e thousands upon thousands of s p e c i e s n a m e s which a p p e a r in t h e l i t e r a t u r e . F u r t h e r m o r e ,
the t a s k i s completely f r u s t r a t i n g because of the g e n e r a l l y inadequate des c r i p t i o n s applied to m o s t of t h e s e n a m e s . The only r e a s o n a b l e solution
s e e m s to be to e s t a b l i s h a new and v e r y r e c e n t s t a r t i n g date f o r t h e a c ceptance of n a m e s of bacteria, a s h a s been suggested many t i m e s in the
p a s t , and to r e q u i r e adequate d e s c r i p t i o n s a s well a s designation and
deposition of type s t r a i n s a s p a r t of the r e q u i r e m e n t s for the valid publication of new n a m e s . Exceptions to the new s t a r t i n g date would be m a d e
for named taxa which a r e adequately d e s c r i b e d and recognizable and for
which c u l t u r e s of type o r neotype s t r a i n s a r e available; f o r o r g a n i s m s
which a r e unique but which cannot be cultivated, a n adequate d e s c r i p t i o n
o r i l l u s t r a t i o n would s e r v e a s the nomenclatural type; the n a m e s of t h e s e
taxa could be r e t a i n e d o r accepted with c r e d i t given t o t h e original a u t h o r s
of the n a m e s . The intent of t h i s action i s to avoid making a u t h o r s spend
t h e i r valuable t i m e s e a r c h i n g through ambiguous l i t e r a t u r e m e r e l y for
the purpose of determining the e a r l i e s t n a m e of the taxon they a r e conc e r n e d with and then, a s in m o s t c a s e s when older synonyms a r e d i s covered, having to c o n s e r v e the l a t e r n a m e o r r e j e c t t h e e a r l i e r n a m e .
,
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JOURNAL
The work involved in updating the d e s c r i p t i o n s and designating type o r
neotype s t r a i n s of t h r e e , four o r even five hundred species i s c o n s i d e r ably l e s s than that entailed in determining the adequacies of thousands
upon thousands of d e s c r i p t i o n s and rejecting thousands upon thousands of
n a m e s a s nomina dubia.
In s u m m a r y , considerable p r o g r e s s h a s been m a d e i n b a c t e r i a l s y s t e m a t i c s , p a r t i c u l a r l y in c h a r a c t e r i z a t i o n and i n approaches t o d e t e r mining t h e relationships of the b a c t e r i a t o e a c h other. N e v e r t h e l e s s t h e
p r e s e n t s t a t e of b a c t e r i a l s y s t e m a t i c s is, frankly speaking, not good,
p r i m a r i l y because much of the older l i t e r a t u r e is inadequate i n t e r m s of
p r e s e n t - d a y knowledge and because type o r neotype s t r a i n s of m o s t of
the named s p e c i e s a r e not available. The d i s o r d e r and confusion that
e x i s t a r e difficult to understand in light of m a n ' s r e m a r k a b l e scientific
achievements, However i n all f a i r n e s s it m u s t be pointed out t h a t m u c h
of s y s t e m a t i c s i s subjective, and it i s the g r e a t difference in human
opinion that is the s o u r c e of much of o u r difficulty. It i s felt, however,
that if c e r t a i n admittedly a r b i t r a r y s t e p s w e r e taken, b a c t e r i a l s y s t e m a t i c s could be placed on a solid foundation and taxonomists could at
l e a s t o p e r a t e on common grounds even if they do not a g r e e with e a c h
other. These steps a r e :
1) r e q u i r e , a s a condition f o r t h e valid publication of the n a m e of a
bacterium, a) that where p r a c t i c a l original d e s c r i p t i o n s of b a c t e r i a contain information on a basic l i s t of c h a r a c t e r i s t i c s ; c e r t a i n c h a r a c t e r i s t i c s
would be r e q u i r e d for a l l bacteria, and additional c h a r a c t e r i s t i c s would
be r e q u i r e d for o r g a n i s m s placed i n specific f a m i l i e s o r g e n e r a , and b)
that a type s t r a i n be designated and that c u l t u r e s of the type s t r a i n be
deposited in a culture collection where the s t r a i n would be available; in
those c a s e s w h e r e the o r g a n i s m cannot be cultivated, t h i s r e q u i r e m e n t
would not pertain,
2) e s t a b l i s h a v e r y r e c e n t date, p e r h a p s even one in t h e future, a s the
beginning date for valid publication of the n a m e s of the b a c t e r i a ; the
p r e s e n t s t a r t i n g date is May 1, 1753; those n a m e s of recognizable taxa
which should be retained because of common usage could be retained
with authorship credited to the o r i g i n a l a u t h o r s , and
3) i n o r d e r t o promote sound p r a c t i c e s in b a c t e r i a l s y s t e m a t i c s , e s tablish, probably through t h e International Committee on Nomenclature
of Bacteria, through national microbiological societies, o r through the
Conference of Biological E d i t o r s , a committee o r committees of qualified
individuals to a s s i s t e d i t o r s of microbiological j o u r n a l s by providing
t h e m with information relating to t h e r e q u i r e m e n t s for the valid publication and legitimacy of n a m e s of b a c t e r i a and t o the nomenclatural-type
concept and p e r h a p s by reviewing p a p e r s on s y s t e m a t i c s submitted f o r
publication.
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