Gram stain

Gram stain (Gram staining or Gram's method), is a met hod of st aining used t o classify
bact erial species int o t wo large groups: gram-posit ive bact eria and gram-negat ive bact eria. It
may also be used t o diagnose a fungal infect ion.[1] The name comes from t he Danish
bact eriologist Hans Christ ian Gram, who developed t he t echnique in 1884.[2]

Micrograph of a gram-positive
coccus and a gram-negative rod.

A Gram stain of mixed

S taphylococcus aureus (S . aureus
ATCC 25923, gram-positive cocci, in
purple) and Escherichia coli (E. coli
ATCC 11775, gram-negative bacilli, in
red), the most common Gram stain
reference bacteria

Gram st aining different iat es bact eria by t he chemical and physical propert ies of t heir cell
walls. Gram-posit ive cells have a t hick layer of pept idoglycan in t he cell wall t hat ret ains t he
primary st ain, cryst al violet . Gram-negat ive cells have a t hinner pept idoglycan layer t hat
allows t he cryst al violet t o wash out on addit ion of et hanol. They are st ained pink or red by
t he count erst ain,[3] commonly safranin or fuchsine. Lugol's iodine solut ion is always added
aft er addit ion of cryst al violet t o st rengt hen t he bonds of t he st ain wit h t he cell membrane.

Gram st aining is almost always t he first st ep in t he ident ificat ion of a bact erial group. While
Gram st aining is a valuable diagnost ic t ool in bot h clinical and research set t ings, not all
bact eria can be definit ively classified by t his t echnique. This gives rise t o gram-variable and
gram-indeterminate groups.

History
The met hod is named aft er it s invent or, t he Danish scient ist Hans Christ ian Gram (1853–
1938), who developed t he t echnique while working wit h Carl Friedländer in t he morgue of t he
cit y hospit al in Berlin in 1884. Gram devised his t echnique not for t he purpose of dist inguishing
one t ype of bact erium from anot her but t o make bact eria more visible in st ained sect ions of
lung t issue.[4] He published his met hod in 1884, and included in his short report t he observat ion
t hat t he t yphus bacillus did not ret ain t he st ain.[5]

Uses

Gram stain of Candida albicans from a

vaginal swab. The small oval
chlamydospores are 2–4 µm in
diameter.

Gram st aining is a bact eriological laborat ory t echnique [6] used t o different iat e bact erial
species int o t wo large groups (gram-posit ive and gram-negat ive) based on t he physical
propert ies of t heir cell walls.[7] Gram st aining can also be used t o diagnose a fungal
infect ion.[1] Gram st aining is not used t o classify archaea, since t hese microorganisms yield
widely varying responses t hat do not follow t heir phylogenet ic groups.[8]

Some organisms are gram-variable (meaning t hey may st ain eit her negat ive or posit ive); some
are not st ained wit h eit her dye used in t he Gram t echnique and are not seen.

Medical
Gram st ains are performed on body fluid or biopsy when infect ion is suspect ed. Gram st ains
yield result s much more quickly t han cult uring, and are especially import ant when infect ion
would make an import ant difference in t he pat ient 's t reat ment and prognosis; examples are
cerebrospinal fluid for meningit is and synovial fluid for sept ic art hrit is.[9][10]

Staining mechanism

Purple-stained gram-positive (left)

and pink-stained gram-negative
(right)

Gram-posit ive bact eria have a t hick mesh-like cell wall made of pept idoglycan (50–90% of
cell envelope), and as a result are st ained purple by cryst al violet , whereas gram-negat ive
bact eria have a t hinner layer (10% of cell envelope), so do not ret ain t he purple st ain and are
count er-st ained pink by safranin. There are four basic st eps of t he Gram st ain:

1. Applying a primary stain (crystal

violet) to a heat-fixed smear of a
 bacterial culture. Heat fixation kills
some bacteria but is mostly used to
affix the bacteria to the slide so that
they do not rinse out during the
staining procedure.
2. The addition of iodine, which binds
to crystal violet and traps it in the
cell
3. Rapid decolorization with ethanol or
acetone
4. Counterstaining with safranin.[11]
Carbol fuchsin is sometimes
substituted for safranin since it
more intensely stains anaerobic
bacteria, but it is less commonly
used as a counterstain.[12]
Summary of Gram stain

Cell color
Application of Reagent
Gram-positive Gram-negative

Primary dye cryst al violet purple purple

mordant iodine purple purple

Decolorizer alcohol/acet one purple colorless

Count er st ain safranin/carbol fuchsin purple pink or red

+ −
Cryst al violet (CV) dissociat es in aqueous solut ions int o CV and chloride (Cl ) ions. These ions
+
penet rat e t he cell wall of bot h gram-posit ive and gram-negat ive cells. The CV ion int eract s
wit h negat ively charged component s of bact erial cells and st ains t he cells purple.[13]

− − +
Iodide (I or I 3) int eract s wit h CV and forms large complexes of cryst al violet and iodine
(CV–I) wit hin t he inner and out er layers of t he cell. Iodine is oft en referred t o as a mordant ,
but is a t rapping agent t hat prevent s t he removal of t he CV–I complex and, t herefore, colors
t he cell.[14]

When a decolorizer such as alcohol or acet one is added, it int eract s wit h t he lipids of t he cell
membrane.[15] A gram-negat ive cell loses it s out er lipopolysaccharide membrane, and t he inner
pept idoglycan layer is left exposed. The CV–I complexes are washed from t he gram-negat ive
cell along wit h t he out er membrane.[16] In cont rast , a gram-posit ive cell becomes dehydrat ed
from an et hanol t reat ment . The large CV–I complexes become t rapped wit hin t he gram-
posit ive cell due t o t he mult ilayered nat ure of it s pept idoglycan.[16] The decolorizat ion st ep is
crit ical and must be t imed correct ly; t he cryst al violet st ain is removed from bot h gram-
posit ive and negat ive cells if t he decolorizing agent is left on t oo long (a mat t er of
seconds).[17]

Aft er decolorizat ion, t he gram-posit ive cell remains purple and t he gram-negat ive cell loses
it s purple color.[17] Count erst ain, which is usually posit ively charged safranin or basic fuchsine,
is applied last t o give decolorized gram-negat ive bact eria a pink or red color.[3][18] Bot h gram-
posit ive bact eria and gram-negat ive bact eria pick up t he count erst ain. The count erst ain,
however, is unseen on gram-posit ive bact eria because of t he darker cryst al violet st ain.
Examples

Gram-positive bacteria

Gram-stain of gram-positive
streptococci surrounded by pus cells

Gram-posit ive bact eria generally have a single membrane (monoderm) surrounded by a t hick
pept idoglycan. This rule is followed by t wo phyla: Bacillot a (except for t he classes
Mollicut es and Negat ivicut es) and t he Act inomycet ot a.[7][19] In cont rast , members of t he
Chloroflexot a (green non-sulfur bact eria) are monoderms but possess a t hin or absent (class
Dehalococcoidet es) pept idoglycan and can st ain negat ive, posit ive or indet erminat e;
members of t he Deinococcot a st ain posit ive but are diderms wit h a t hick pept idoglycan.[7][19]

Hist orically, t he gram-posit ive forms made up t he phylum Firmicut es, a name now used for t he
largest group. It includes many well-known genera such as Lactobacillus, Bacillus , Listeria ,
Staphylococcus , Streptococcus , Enterococcus , and Clostridium.[20] It has also been
expanded t o include t he Mollicut es, bact eria such as Mycoplasma and Thermoplasma t hat
lack cell walls and so cannot be Gram-st ained, but are derived from such forms.[21]

Some bact eria have cell walls which are part icularly adept at ret aining st ains. These will
appear posit ive by Gram st ain even t hough t hey are not closely relat ed t o ot her gram-posit ive
bact eria. These are called acid-fast bact eria, and can only be different iat ed from ot her gram-
posit ive bact eria by special st aining procedures.[22]
Gram-negative bacteria

Gram negative Neisseria gonorrhoeae

and pus cells

Gram-negat ive bact eria generally possess a t hin layer of pept idoglycan bet ween t wo
membranes (diderm).[23] Lipopolysaccharide (LPS) is t he most abundant ant igen on t he cell
surface of most gram-negat ive bact eria, cont ribut ing up t o 80% of t he out er membrane of E.
coli and Salmonella .[24] Most bact erial phyla are gram-negat ive, including t he cyanobact eria,
green sulfur bact eria, and most Pseudomonadot a (except ions being some members of t he
Ricket t siales and t he insect -endosymbiont s of t he Ent erobact eriales).[7][19]

Gram-variable and gram-

indeterminate bacteria
Some bact eria, aft er st aining wit h t he Gram st ain, yield a gram-variable pat t ern: a mix of pink
and purple cells are seen.[16][25] In cult ures of Bacillus, Butyrivibrio, and Clostridium, a
decrease in pept idoglycan t hickness during growt h coincides wit h an increase in t he number
of cells t hat st ain gram-negat ive.[25] In addit ion, in all bact eria st ained using t he Gram st ain,
t he age of t he cult ure may influence t he result s of t he st ain.[25]

Gram-indet erminat e bact eria do not respond predict ably t o Gram st aining and, t herefore,
cannot be det ermined as eit her gram-posit ive or gram-negat ive. Examples include many
species of Mycobacterium, including Mycobacterium bovis , Mycobacterium leprae and
Mycobacterium tuberculosis , t he lat t er t wo of which are t he causat ive agent s of leprosy and
t uberculosis, respect ively.[26][27] Bact eria of t he genus Mycoplasma lack a cell wall around
t heir cell membranes,[9] which means t hey do not st ain by Gram's met hod and are resist ant t o
t he ant ibiot ics t hat t arget cell wall synt hesis.[28][29]

Orthographic note
The t erm Gram staining is derived from t he surname of Hans Christ ian Gram; t he eponym
(Gram) is t herefore capit alized but not t he common noun (st ain) as is usual for scient ific
t erms.[30] The init ial let t ers of gram-positive and gram-negative, which are eponymous
adject ives, can be eit her capit al G or lowercase g, depending on what st yle guide (if any)
governs t he document being writ t en. Lowercase st yle is used by t he US Cent ers for Disease
Cont rol and Prevent ion and ot her st yle regimens such as t he AMA st yle.[31] Dict ionaries may
use lowercase,[32][33] uppercase,[34][35][36][37] or bot h.[38][39] Uppercase Gram-positive or Gram-
negative usage is also common in many scient ific journal art icles and publicat ions.[39][40][41]
When art icles are submit t ed t o journals, each journal may or may not apply house st yle t o t he
post print version. Preprint versions cont ain whichever st yle t he aut hor happened t o use. Even
st yle regimens t hat use lowercase for t he adject ives gram-positive and gram-negative st ill
t ypically use capit al for Gram stain.

See also

Bacterial cell structure

Ziehl–Neelsen stain

References

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