FEMS Microbiology Letters Advance Access published February 29, 2016

MiniReview:

Diversity of Phage Infection Types and Associated Terminology:

The Problem with “Lytic or Lysogenic”

Zack Hobbs1 and Stephen T. Abedon2

1
EpiBiome, Inc.; South San Francisco, California, USA
2
Department of Microbiology, The Ohio State University; Mansfield, Ohio USA

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Abstract:

Bacteriophages, or phages, are viruses of the members of domain Bacteria. These viruses play numerous
roles in shaping the diversity of microbial communities, with impact differing depending on what
infection strategies specific phages employ. From an applied perspective especially are communities
containing undesired or pathogenic bacteria that can be modified through phage-mediated bacterial
biocontrol, that is, through phage therapy. Here we seek to categorize phages in terms of their infection
strategies as well as review or suggest more descriptive, accurate, or distinguishing terminology.
Categories can be differentiated in terms of (1) whether or not virion release occurs (productive
infections vs. lysogeny, pseudolysogeny, and/or the phage carrier state), (2) the means of virion release
(lytic vs. chronic release), and (3) the degree to which phages are genetically equipped to display
lysogenic cycles (temperate vs. non-temperate phages). We address in particular the use or over-use of
what can be a somewhat equivocal phrase, “Lytic or lysogenic”, especially when employed as a means of
distinguishing among phages types. We suggest that the implied dichotomy is inconsistent with both
modern as well as historical understanding of phage biology. We consider, therefore, less ambiguous
terminology for distinguishing between “Lytic” versus “Lysogenic” phage types.

Keywords: Lytic phage, Obligately lytic, Phage therapy, Professionally lytic, Strictly lytic, Temperate
phage

Running title: Phage Infection Types and Associated Terminology


Correspondence to: Stephen T. Abedon; Department of Microbiology, The Ohio State University, 1680 University
Dr., Mansfield, OH 44906 USA; Tel.: 419.755.4343; Fax: 419.755.4327; Email: abedon.1@osu.edu.

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One-sentence summary: The phrase “Lytic or lysogenic” we suggest can be problematic as most phages
which display lysogeny also are “Lytic” while bacteria are lysogenic, not phages.

Introduction:

In this MiniReview we consider the diversity of phage types as based on potential infection
strategies, particularly productive or lysogenic along with lytic release versus chronic release, with
emphasis on what major variants should be called (see Table 1 for glossary of terms). In a few of these
cases, efforts towards naming do not appear to have been a priority within the community. In other
cases, perhaps too many names exist, at least some of which are less precise than may be desirable. We
place particular emphasis on addressing what we will argue is common misuse of the phrase, “Lytic or
lysogenic”, as a means of distinguishing non-temperate from temperate phages. We begin, however,

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with a scheme for classifying phages, in terms of infection strategies, into four distinct categories: (I)
lytic and non-temperate, (II) chronic and non-temperate, (III) lytic and temperate, or (IV) chronic and
temperate.

Strategies of Phage Infection and Release

Phages can be defined as semi-autonomous genetic elements that at some point in their life
cycles exist as encapsidated genomes, particularly infectious capsids that are not found within the
bounds of host cells. Most viruses can be defined similarly, though especially fungal viruses
(mycoviruses) do not appear to exist as host-unassociated virions or, in a few instances, even as
encapsidated genomes (Hyman and Abedon 2012). Here, however, our emphasis primarily is on
bacteriophages.

Phages, in terms of their encapsidation and location, can exist in three possible states: (1)
intracellularly and unencapsidated, (2) intracellularly and packaged within mature virions, or (3) both
encapsidated and extracellular. The first state can be further subdivided into what can be described as a
“Vegetative phase” (Lwoff 1953) or productive cycle versus existing as a prophage, that is, displaying a
lysogenic cycle. The second state can be distinguished from phage genomes that instead are not
packaged until the virion release step. The third state consists of free phages, that is, assembled virions
that are no longer found within their bacterial host. Abbreviations, used here-forward to describe these
different states include V for Vegetative phase, P for Prophage, B for genome packaging into mature
virions Before release, D for genome packaging into mature virions During release, and F for Free
phages.

To distinguish among phages, but intentionally without using more standard terms as otherwise
are under discussion, we categorize phages into four distinct types using the abbreviations introduced at
the end of the previous paragraph. These are

I. “V  B  F” phages

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 II. “V  D  F” phages
III. “V or P  B  F” phages
IV. “V or P  D  F” phages

The phrase “V or P” is intended to imply that these phages can display either V- or, instead, P- and then
V-type infection states, i.e. Vegetative or Prophage states (Table 1). Productive infections, ones which
produce Free phage virions (“F”), can be contrasted with infections that instead may be described as
“Reductive” (Lwoff 1953), particularly lysogenic infections (“P”) but also, though not otherwise
considered here, pseudolysogenic ones (Miller and Day 2008; Abedon et al. 2009; Abedon 2009; Los and
Wegrzyn 2012).

In more familiar terms, the phage types described above and in Figure 1 thus include the
following: (I) lytic phages that do not display lysogenic cycles (“V  B  F” phages), (II) chronically
releasing phages that do not display lysogenic cycles (“V  D  F”), (III) lytic phages that can display

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lysogenic cycles (i.e. temperate phages; “V or P  B  F”), and (IV) chronically released phages also
that can display lysogenic cycles (which are temperate phages as well; “V or P  D  F”). In other
words, we can contrast phage types into lytic versus chronic (“B” versus “D” phages) as well as not-
temperate versus temperate (“V-only” versus “V or P” phages). See Table 1 and Table 2 for further
consideration.

Example phages

Examples of obligately lytic phages (category I) include Max Delbrück’s well-studied tailed
phages T1-T7 (Abedon 2000), with phage T4 the most thoroughly studied (Karam 1994; Miller et al.
2003). While tailed phages are always lytic, they are not always obligately lytic (i.e. see category III,
below). RNA-genomed phages of families Leviviridae (van Duin and Tsareva 2006) and Cystoviridae
(Mindich 2006) we can speculate are consistently obligately lytic. The Microviridae (Fane et al. 2006),
however, may include members which are not obligately lytic (Krupovic and Forterre 2011). Note that
there exists as well variation on how lysis is induced by various lytic phages (Young 2014).

Phage M13 is one of the best studied category II phages (non-temperate, chronic) (Russel and
Model 2006). As with lysis, the mechanisms by which chronic release occurs varies among phages, i.e.
via extrusion as seen with phage family Inoviridae (Russel and Model 2006) versus via budding as seen
with phage family Plasmaviridae (Maniloff and Dybvig 2006).

Phage λ is the most intensely studied of the tailed, temperate phages (Hendrix et al. 1983;
Ptashne 2004; Casjens and Hendrix 2015) and thereby is representative of category III.

Phage CTXΦ (Mai-Prochnow et al. 2015), which encodes the cholera toxin, is the most
prominent example of category IV, though with additional members of phage family Inoviridae also
thought to be temperate (Yamada et al. 2007). In addition, mycoplasma phage L2, a member of phage

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family Plasmaviridae (Maniloff and Dybvig 2006), has been described as temperate (Maniloff et al.
1994).

See Abedon (2008) and Abedon et al. (2009) for additional discussion of phage types
distinguished by infection as well as Calendar and Abedon (2006) for numerous individual reviews of
specific phage types. See Table 1 for association of these phage types with specific terms and concepts
as considered in this MiniReview.

“Lytic or Lysogenic”

Use of the phrase, “Lytic or lysogenic”, as a means of distinguishing among phage types, may be
becoming increasingly common (e.g., see the following paragraph for its use in publications). This we
believe is of concern particularly during discussions of the medical procedure known as a phage therapy,

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e.g. (Kutter et al. 2010; Abedon et al. 2011; Abedon 2015), which as typically practiced tends to strive to
avoid using phages that can be categorized into types II, III, or IV, i.e. phages that are not obligately lytic.
In this section we therefore strive to illuminate the problems associated with the dichotomy of “Lytic or
lysogenic”, particularly as a means of distinguishing among phage types (“Phage” or “Phages”, etc., in
the searches below), even if the intended meaning is not altogether lost.

Using Google Scholar, a search on "lytic or lysogenic phage" OR "lytic or lysogenic phages" OR
"lytic or lysogenic bacteriophage" OR "lytic or lysogenic bacteriophages" (with quotation marks) yielded
36 results. The dates of these results are 2015 (4 results), 2014 (6), 2013 (6), 2012 (4), 2011 (4), 2010 (1),
2009 (2), 2007 (3), 2005 (2), 2002 (1), 1989 (1), 1984 (1), and 1967 (1), that is, most of this usage is fairly
recent. At least one of these results, in 2015, refers, however, to a “lytic or lysogenic phage life cycle”
(emphasis added). By comparison, a Google Scholar search on "lytic or lysogenic cycle" OR "lytic or
lysogenic cycles" yields a total of 101 results. By decades, numbers are 2006-2015 (65 results), 1996-
2005 (23), 1986-1995 (6), and then two additional results from 1968. Both phrases thus appear to be
becoming increasingly part of the phage biology vernacular.

Part of the problem with use of the phrase, “Lytic or lysogenic”, is that bacteria can be lysogenic
while phages instead can be temperate, but not vice versa (see Figure 2 for an exploration of the history
of the usage of “Lysogenic phage” versus “Temperate phage”). Thus, from Lwoff (1953), p. 271
(emphasis his): “Lysogeny is the hereditary power to produce bacteriophage. A lysogenic bacterium is a
bacterium possessing and transmitting the power to produce bacteriophage. Each bacterium of a
lysogenic strain gives rise to a lysogenic clone…” Also from Lwoff (1953), p. 273 (emphasis his): “The
power to lysogenize is the property of temperate phages…” The vast majority of temperate phages in
addition display lytic cycles when productively infecting, that is, rather than chronically releasing virion
progeny, hence the term, “Lysogenic”, to describe them. What, then, is meant when contrasting “Lytic”
with “Lysogenic”?

Though most temperate phages during productive cycles release virions via lysis – that is, during
lytic cycles – nonetheless in all likelihood the intended use of “Lytic” in “Lytic or lysogenic” as

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descriptions of phage types is as shorthand for the “V  B  F” (I) phages, that is, lytic, non-temperate
phages. The term virulent, however, is often employed preferentially to “Lytic, non-temperate” (Lwoff
1953), thus, “Virulent or lysogenic” might be employed rather than “Lytic or lysogenic”. Unfortunately,
the term virulent has had multiple meanings over the years with regard to phages (next section). To
avoid this issue, it can be preferable, for example, to use the phrase “Obligately lytic” instead of
“Virulent” to describe lytic, non-temperate phages. As an aside, note that “Obligatorily lytic” (in
quotation marks) yields 26 Google Scholar results versus 142 for “Obligately lytic”.

There exist additional terms that can be used equivalently to obligately lytic. These include
“Strictly lytic” (which in a within-quotes Google Scholar search yields 223 results), “Obligate lytic” (113
Google Scholar results), “Professionally lytic” (10 results, e.g. Brüssow and Kutter 2005; Kutter 2008;
Henein 2013; Kutter et al. 2013; Miroshnikov et al. 2014), “Professionally virulent" (Górski et al. 2009),
and “Professional virulent” (9 results, e.g. Brüssow and Kutter 2005; Brüssow 2005; Brüssow 2007;
Weiss et al. 2009; Letarov and Kulikov 2009; Letarov et al. 2010; Sarker et al. 2012; Bourdin et al. 2014).

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All of these terms are preferable to simply “Lytic” when describing phages, unless one is intentionally
referring simultaneously to category I phages (“V  B  F”, i.e. strictly lytic) and category III phages (“V
or P  B  F”, i.e. temperate phages that can display both lytic and lysogenic cycles). That is, the
phrase “Lytic phage” (with 6,080 Google Scholar results for a “Lytic phage” OR “Lytic phages” search)
distinguishes among phages particularly in terms of their mode of virion release, lytic rather than
chronic, rather than in terms of their ability to display lysogenic cycles. These various phage descriptors
and others are summarized in Table 1. (For completeness, note that a “Chronic phage” OR “Chronic
phages” search yields 49 Google Scholar results.)

Further Arguments against “Virulent”

The term “Virulence” can be used as a description of those phages that are particularly adept at
inflicting substantial destruction on their host (Smith et al. 1987). Within an historical context, this is as
occurs primarily to bacterial cultures rather than specifically to individual bacterial cells (Summers 1991).
Phages that are less adept at lysing bacterial cultures therefore can be described as being less virulent.
These can include both temperate phages and also chronically releasing phages, though the latter are
not an emphasis of this section. In the case of temperate phages, this lower virulence occurs because a
subset of individual phage infections display lysogenic rather than lytic cycles. The resulting bacterial
lysogens also tend to be insensitive to the phage types that are lysogenically infecting them – due to
display of superinfection immunity (Hershey and Dove 1983; Fogg et al. 2010; Hyman and Abedon 2010;
Casjens and Hendrix 2015) – and therefore are inclined to resist phage-mediated clearance.

A further complication on the term, “Virulent”, is that otherwise temperate phages that are
genetically defective in their ability to lysogenize are described as virulent mutants (Lwoff 1953) (2240
Google Scholar results for "virulent mutant" OR "virulent mutants"; though refining the search to
include the term “phage”, not in quotes, yields only 1140 results). Virulence, of course, began as a term
that was not specific to phage biology and which, according to the Oxford Dictionary, dates back at least

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to the 1800s in the English language. “Virulent” was then legitimately coopted close to the beginning of
the study of phage biology to describe phages that in a traditional sense are indeed more virulent, at
least against bacterial cultures. The term then came to be used to a large extent as a description of
phages that do not display lysogenic cycles and which otherwise are lytic, including as referring to
virulent mutants of otherwise temperate phages (Ptashne 2004).

The term “Obligately lytic”, or “Strictly lytic”, etc., contrasting the ambiguity of “Virulent”, is
meant to explicitly imply an unavoidable display of lytic, productive cycles upon successful infection
(Abedon 2008). The term “Obligately productive”, though almost never used (no PubMed hits, and no
Google Scholar hits that are not associated with one of us), would imply a similarly requisite display of
either lytic or chronic infections, depending on the phage, rather than referring exclusively to lytic
infections, as is implied by “Obligately lytic”. Consider also “Obligately chronic” (Table 2). All of these
terms can be viewed as more-precise alternatives to the concept of “Virulent” to describe phages that
are not temperate. In addition, note that obligately lytic phages may or may not be recent descendants

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of temperate phages and thus may or may not represent virulent mutants.

Within this framework, what then is meant by “Lytic or lysogenic”? This question is particularly
relevant within the context of phage therapy where “Lytic” phages typically are considered to be
preferable to “Lysogenic” ones. One simple answer could be that “Lytic” is being used equivalently to
“Obligately lytic” or “Strictly lytic” as a substitute for “Virulent”. Such usage can be problematic,
however, since with phage therapy a temperate phage ancestry can be of concern, even if lysogenic
cycles themselves do not occur, since temperate phages especially have been shown to carry bacterial
virulence factor genes (Hyman and Abedon 2008; Kuhl et al. 2012; Christie et al. 2012). In response to
this concern, one could replace “Lytic” – in “Lytic or lysogenic” – with the phrase, “Virulent meaning
obligately lytic but nevertheless not a virulent mutant of a temperate phage”. Obviously, however, such
phrasing is too complex to become a part of everyday practice. Instead, and perhaps ideally, one might,
for example, use the otherwise somewhat underutilized “Professionally lytic” to specifically describe
those phages that are both obligately lytic and not closely related to temperate phages, that is, lytic
phages that are both not temperate and not virulent mutants of temperate phages (Curtright and
Abedon 2011). This perspective we summarize in Figure 3.

“Lytic and Lysogenic”

As alternatives to “Lytic or lysogenic”, we thus can have “Non-temperate or temperate”,

“Virulent or temperate”, “Obligately lytic or temperate”, “Strictly lytic or temperate”, “Virulent mutant
or temperate”, and/or “Professionally lytic or temperate”. “Lytic or temperate”, by contrast, does not
really work, despite the increasing popularity of “Lytic or lysogenic” as a description of phage types,
since most temperate phages, as noted, are also lytic phages. Nonetheless, what all of these alternative
phrases have in common is their use of the term, “Temperate”, and this is rather than “Lysogenic”. What
is the justification for this substitution?

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 “Lysogenic” literally means “Generating lysis” (Lwoff 1953); see Lwoff for discussion of the early
history of the study of lysogeny. The majority of phages, however, appear to be lysis generating. Indeed,
that is precisely what should be meant by the term “Lytic phage”—a lytic phage literally is a lysis-
generating phage. As a consequence, it logically can be preferable to state “Lytic and lysogenic” rather
than “Lytic or lysogenic”. The apparent equivalence of these two terms – terms that in “Lytic or
lysogenic” seemingly are used to imply polar opposites – stems from an error in usage. That is, rather
than a description of a phage property, the term “lysogenic” when correctly employed is a description of
a bacterial property. Specifically, certain bacterial cultures exist that when added to cultures of certain
other bacterial strains can give rise to the lysis of these other cultures. The first culture thus has the
property of being able to generate lysis in the second culture. This first culture therefore is lysis
generating, that is, lysogenic (Lwoff 1953).

As scientific understanding has advanced, at least one of the conditions associated with this
lysogenic property, that it can be a property of bacterial cultures, came to be associated with what

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today we refer to as lysogens: bacteria that harbor prophages. As part of this transition in meaning, note
that Lwoff (1953), in his classic review, does not appear to even use “Lysogen” but instead refers to a
“Lysogenic bacterium”, as too does Adams (1959), though “Lysogen” is found in Stent (1963). The term
“Temperate phage”, by contrast, does appear in Lwoff as does also “Temperate” alone, as in, “The
temperate or virulent character of a phage” (p. 326).

“Professionally Lytic or Temperate”

With the phrase “Lytic or lysogenic”, as considered here, it is probable that virulent, obligately
lytic, or strictly lytic is what generally is intended by “Lytic”, while “Temperate” without question should
replace “Lysogenic”. “Obligately lytic or temperate” (4 Google Scholar results), “Strictly lytic or
temperate” (2 results), or instead “Virulent or temperate” (136 results), therefore could be legitimate
alternatives to “Lytic or lysogenic”. “Virulent”, though, and as noted, is burdened not only with historical
ambiguity but also may not be the most ideal of descriptors in terms of the marketing of virus-based
medicinals. Indeed, within a phage therapy context, “Professionally lytic or temperate” (zero results in
either Google Scholar or Google Books) would seem to be the most meaningful alternative to the highly
questionable, “Lytic or lysogenic”, that is, so long as “Professionally lytic” phages contrast not just with
temperate phages but with virulent derivatives of temperate phages as well (Table 2; Figure 3). We
suggest therefore that an effort might be made to replace even informal use of “Lytic or lysogenic” with
“Professionally lytic or temperate”, assuming, of course, that this is the distinction which is intended.

Conclusion

In this MiniReview we have considered the diversity of phage types from the perspective of
productive versus lysogenic infections along with lytic versus chronic release. We have emphasized
issues associated with use of the terms “Lytic” and “Lysogenic” as stand-alone descriptors of phages and

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their properties, which either can be ambiguous (as is the case for “Lytic”) or etymologically
inappropriate (for “Lysogenic” as a descriptor of phages). In particular, we have criticized the phrasing,
“Lytic or lysogenic”. We note as well that “Lysogenic”, logically, should never be used in association with
non-lytic viral release from cells, that is, to describe phages that release their virions chronically or
continuously from infected bacteria (category II). “Lysogenic”, strictly speaking, also is not an accurate
general descriptor of all phage latent infections since “Lysogens” associated with non-lytic viruses
(category IV), such as the filamentous phage, CTXΦ (Mai-Prochnow et al. 2015), are not lysis generating.

Despite our numerous concerns with use of the phrase, “Lytic or lysogenic”, there nonetheless
exist circumstances in which the phrase unquestionably is valid. This is seen when describing the initial
infection of a bacterium by a lytic, temperate phage (category III), such as phage λ, where the choice is
between a lytic cycle or, instead, a lysogenic cycle, thus, “Lytic or lysogenic [cycles]”. The suitability of
“Lytic or lysogenic” as shorthand for distinguishing among phage infection types – particularly lytic
versus lysogenic cycles as one sees with phage λ – does not, however, imply an equivalent legitimacy of

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extension of this phrasing to distinguish among phage types.

Acknowledgements

STA has consulted and served on advisory boards for companies with phage therapy interests
and maintains the websites phage.org and phage-therapy.org. ZH is an employee of EpiBiome Inc, a
phage-based biotech company. Neither received help or financial support in writing the manuscript.

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Table 1: Glossary of Terms.

1 2 3
Term States Phages Category Definition as Employed Here
Mutated temperate phage that as a consequence
λ
Clear mutant I is unable to display lysogenic cycles and thus
mutant
forms clear rather than turbid plaques
Productive infections in which virions are released
M13,
Chronic D II, IV over long intervals without substantial disruption
CTXΦ
of host cells
M13, Phages whose productive infections are chronic
Chronic phage D II, IV
CTXΦ and which may nor may not be temperate
Chronic phage that is able to display lysogenic
Chronic temperate phage P,V,D CTXΦ IV cycles, i.e. during which virions are neither

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Encapsidated during release
Encapsidated intracellularly
Free phage
Induction
Latent infection
Lysogenic cycle
Lysogenic infection
Lysogen
Lysogenic (current def.)
Lysogenic (historical def.)
produced nor released
M13, Referring to virion maturation during release, e.g.
D II, IV
CTXΦ as seen with filamentous phages
Referring to virion maturation prior to release,
B T4, λ I, III
e.g. as seen with tailed phages
F all All Mature virion that is extracellularly located
Intracellular transition of a lysogenic infection to a
V λ, CTXΦ III, IV
productive infection
For bacteriophages, these predominantly are
P λ, CTXΦ III, IV
lysogenic infections
Phage replication as a prophage that does not
P λ, CTXΦ III, IV
directly result in virion production or release
P λ, CTXΦ III, IV Ongoing phage existence as a prophage
Bacterium that is latently infected with a
P λ, CTXΦ III, IV
temperate phage
Property of a bacterium indicating that it is
P λ, CTXΦ III, IV
hosting a prophage/lysogenic cycle
Property of a bacterium indicating an ability to
λ III release factors that are able to lyse other bacterial
cultures

1
“States” refers to distinguishing characteristics as abbreviated throughout the manuscript: “B” refers to phage
genome packaging and virion maturation that occurs Before virion release (property of lytic phages only), “D”
refers to phage genome packaging and virion maturation that occurs During virion release (property of chronic
phages only), “F” refers to Free phages, “P” refers to existence as a Prophage (a property only of temperate
phages), “V” refers to display of a Vegetative phase. Vegetative phases are a property of all productive phage
infections, though to avoid clutter this is mostly not explicitly indicated in the table. Either B or D states also are
properties of all productive phage infections as too, post infection, is the F state. The sequence “P,V” indicates the
existence of either Prophage or Vegetative states. P and V will not occur simultaneously during the same infection
though can occur sequentially, particularly P  V, which then will proceed through B or D states depending on the
phage, culminating in all cases, post successful productive infection, in an F state.
2
Example phages are as discussed in the main text.
3
Categories are “I” meaning lytic and non-temperate, “II” meaning chronic and non-temperate, “III” meaning lytic
and temperate, and “IV” meaning chronic and temperate.

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Lysogenic (literal def.) T4, λ I, III Ability to generate lysis in bacterial cultures
Lysogeny P λ, CTXΦ III, IV A prophage-containing latent phage infection
Productive infections in which phage virions are
Lytic B T4, λ I, III released over short intervals with lethal disruption
of host cells
Lytic cycle B T4, λ I, III Vegetative phage replication that ends with lysis
Productive infection in which virions are released
Lytic infection B T4, λ I, III
via host-cell lysis
Phages whose productive infections are lytic (may
Lytic phage B T4, λ I, III
nor may not be temperate phages)
Lytic temperate phage P,V,B λ III Lytic phage that is able to display lysogenic cycles
Description of a phage that is unable to display
Non-temperate V T4, M13 I, II
lysogenic cycles
Phage that upon infection is inherently unable to
Obligately lytic B T4 I
display lysogenic cycles or chronic release

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Phage that upon infection is inherently unable to
Obligately productive B T4, M13 I, II
display lysogenic cycles
Vegetative phage replication associated with
Productive cycle V all all
either chronic or lytic release
Product of either adsorption or induction
Productive infection V all all involving vegetative state and subsequent phage
release
Phage that is both obligately lytic and not recently
Professionally lytic B T4 I
descended from a temperate ancestor
Prophage P λ, CTXΦ III, IV Phage genome as it exists during lysogenic cycles
Latent infection during which neither phage
genome replication nor prophage formation
Pseudolysogeny occurs; other definitions exist and in some cases
the phrase ‘carrier state’ has been used
synonymously
Strictly lytic B T4 I Used synonymously with obligately lytic
Description of a phage that is able to display
Temperate P,V λ, CTXΦ III, IV
lysogenic cycles
An intracellular phage genome that has not been
Unencapsidated all all
packaged into a virion particle
Replicating phage genome that is not a prophage
Vegetative V all all
(contrast both lysogenic and pseudolysogenic)
Description of ability of phage populations to lyse
Virulence B
cultures of bacterial hosts
Virulent B T4 I Often used synonymously with obligately lytic
Clear temperate phage mutant that can form
λ
Virulent mutant B I plaques even on lysogens formed by the phage
mutant
wild-type parent

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Table 2: Distinguishing among Different Phage Types in Terms of Basic Phage Infection Properties.4

Phage Types (below; characteristics to right) V5 P6 V or P  V7 B8 D9 F10

All phages (I, II, III, or IV) +     +
Obligately Productive (I or II)11 +     +
Chronic (II or IV) +    + +
Obligately Chronic (II)12 +    + +
Lytic (I or III) +   +  +
Obligately or Strictly Lytic (I) +   +  +
Professionally Lytic (I) +   +  +
Virulent, meaning Obligately Lytic (I) +   +  +
Virulent, meaning Highly Destructive of Cultures (I)13 +   +  +

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Virulent, meaning Mutant of a Temperate Phage (I) +  + +  +
Temperate (III or IV) + + x   +

4
“+” implies always a characteristic of, “” implies never a characteristic of, and “” implies may or may not be a
characteristic of.
5
Can exist in a Vegetative state, which is a reproductive, non-prophage, bacterium-infecting, phage genome.
6
Can exist as a Prophage, at least for a given host and conditions; distinguishes temperate (+) from non-temperate
() phages.
7
Mutant and/or descendant of temperate phage, one that no longer can display lysogenic cycles; distinguishes
obligately productive descendants of temperate phages (+) from either obligately productive phages that are not
descendants of temperate phages () or that simply are temperate phages themselves and therefore not obligately
productive (x).
8
Exists as a mature virion prior to release from host bacteria; distinguishes between chronic () from lytic (+)
phages.
9
Does not exist as a mature virion prior to release from host bacteria; distinguishes between chronic (+) from lytic
() phages.
10
Exists as a free phage at some point in life cycle.
11
Not a commonly used descriptor.
12
Not a commonly used descriptor.
13
In this case the culture in question is assumed to consist exclusively of bacteria that do not support lysogenic
infections by the phage in question, even if that phage is capable of displaying lysogenic infections when infecting
different bacterial strains.

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Figure 1

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