ARTICLE IN PRESS

FOOD
MICROBIOLOGY
Food Microbiology 24 (2007) 703710
www.elsevier.com/locate/fm

Combined effect of packaging atmosphere and storage temperature on

growth of Listeria monocytogenes on ready-to-eat shrimp
Thomas J. Rutherforda, Douglas L. Marshalla,, Linda S. Andrewsb,
Patti C. Cogginsa, M. Wes Schillinga, Patrick Gerardc
a
Department of Food Science, Nutrition, and Health Promotion, Mississippi Agricultural and Forestry Experiment Station,
Mississippi State University, Box 9805 Mississippi State, MS 39762-9805, USA
b
Experimental Seafood Processing Laboratory, Coastal Research and Extension Center,
Mississippi State University, 3411 Frederick St., Pascagoula, MS 39567, USA
c
Experimental Statistics Unit, Mississippi Agricultural and Forestry Experiment Station,
Mississippi State University, Box 9653, Mississippi State, MS 39762-9745, USA

Received 25 January 2007; received in revised form 27 March 2007; accepted 28 March 2007
Available online 5 April 2007

Abstract

Cooked, peeled, and deveined shrimp were inoculated with a 5 strain mixture of Listeria monocytogenes and packaged in air, vacuum,
and a 100% carbon dioxide modied atmosphere. The packaged shrimp were then stored at 3, 7, and 12 1C for 15 days to monitor the
growth of L. monocytogenes and psychrotrophic bacteria. Uninoculated shrimp were also subjected to sensory evaluation by a trained
panel to measure odor and appearance over the storage period. Results demonstrated that shrimp packaged in CO2 and stored at 3 1C
did not permit growth of L. monocytogenes during the 15-day storage period, while all other packaging/temperature combinations
allowed for multiplication of the bacterium. Carbon dioxide packaging also resulted in the slowest growth of psychrotrophic bacteria and
resulted in shrimp having acceptable sensory odor and appearance scores at the end of storage. When strict temperature control is
difcult, such as during processing, transportation, retail display, or home use, additional antimicrobial hurdles may be necessary to
ensure safety.
r 2007 Elsevier Ltd. All rights reserved.

Keywords: Shrimp safety; Listeria monocytogenes; MAP; Sensory quality

1. Introduction When developing new products that promote shrimp as

The harvesting of shrimp holds great economic impor-
tance for the U.S. Gulf of Mexico region, amounting to
72% of the total U.S. catch (NOAA, 2003). However,
this domestic supply does not meet increasing consumer
demand for shrimp. As a result, much of the shrimp sold
in the U.S. comes from foreign markets (NOAA, 2003).
In addition to the sheer number of imports avai-
lable, imported shrimp can usually be obtained for
lower prices than domestic shrimp. This puts added
pressure on domestic processors to develop new value-
added products.
Corresponding author. Tel.: +1 662 325 8722; fax: +1 662 325 8728.
E-mail address: microman@ra.msstate.edu (D.L. Marshall).
the primary ingredient, the product developer has two
main concerns, product safety and shelife. It is widely
accepted that fresh shrimp are highly perishable (Bazemore
et al., 2003), thus the predominant market form is as frozen
raw or cooked products. A refrigerated ready-to-eat (RTE)
product would be more convenient for the consumer.
Unfortunately, even frozen RTE shrimp can harbor human
pathogens (Duran and Marshall, 2005). In addition to
enteric pathogens, Listeria monocytogenes is also of
concern for RTE seafoods (Farber, 1991b; Dillon and
Patel, 1992; Gecan et al., 1994; Shineman and Harrison,
1994; Valdimarsson et al., 1998; Notermans and Hoorn-
stra, 2000; Hathed et al., 2003; Suwansonthichai and
Rengpipat, 2003), with an occurrence rate of up to 11% in
raw Gulf Coast shrimp (Motes, 1991; Gecan et al., 1994).

0740-0020/$ - see front matter r 2007 Elsevier Ltd. All rights reserved.
doi:10.1016/j.fm.2007.03.011
 ARTICLE IN PRESS
704 T.J. Rutherford et al. / Food Microbiology 24 (2007) 703710
Several properties make L. monocytogenes a concern in
refrigerated RTE shrimp (Buchanan, 1991; Gecan et al.,
1994; Lionberg et al., 2003). These properties include: (1)
L. monocytogenes is commonly found in shrimp and other
seafood products; (2) it is routinely found in processing
environments as a post-processing contaminant; (3) it has
the ability to multiply at refrigerator temperatures; and (4)
it possesses the ability to reach infectious doses without
spoiling foods. These factors along with the bacteriums
ability to grow in the presence of salt and reduced oxygen
atmospheres makes control of L. monocytogenes a formid-
able problem in RTE seafoods (Dorsa et al., 1993;
Fernandez et al., 1997; Marshall, 2004; Mejlholm et al.,
2005).
Many methods of packaging and storage have been
investigated to control the proliferation of pathogens and
spoilage bacteria in RTE foods. Modied atmosphere
packaging (MAP) entails removal of air from a container
followed by reintroduction of an altered or modied
atmosphere. This new atmosphere may contain carbon
dioxide, nitrogen, and/or oxygen (Genigeorgis, 1985;
Daniels et al., 1985; Farber, 1991a). The actual percentage
of each gas in the mixture is dependent upon the
application; however, CO2 is almost always present in the
greatest quantity because it is effective at extending shelife
by increasing the lag phase and the generation time of most
aerobic spoilage microorganisms (Lannelongue et al., 1982;
Layrisse and Matches, 1984; Matches and Layrisse, 1985;
Marshall et al., 1991; Bak et al., 1999). Reduced oxygen
atmospheres may control the growth of aerobic spoilage
bacteria; however, the same environment may be benecial
for proliferation of psychrotrophic, facultatively anaerobic
or strictly anaerobic pathogens, such as nonproteolytic
Clostridium botulinum and L. monocytogenes in seafoods
(Dorsa and Marshall, 1995; Pothuri et al., 1995; Gimenez
et al., 2002; Mejlholm et al., 2005). It is well known,
however, that high CO2 MAP can reduce the growth rate
of L. monocytogenes relative to growth in air or under
vacuum (Marshall et al., 1991, 1992; Oh and Marshall,
1995; Pothuri et al., 1995).
Another major area of focus in regard to a refrigerated
RTE shrimp product is shelife. Two factors contribute to
quality degradation in stored shrimp. The rst defect,
melanosis, is a black discoloring on the surface of shrimp
that is caused by the activity of polyphenoloxidase, and
occurs only in raw shell-on shrimp (Benner et al., 1994).
Another group of signicant enzymatic reactions result in
the production of protein breakdown products such as
ammonia, indole, methanethiol, putrescine, trimethyl
amine and other off-odor compounds that are caused by
the growth of spoilage bacteria (Chang et al., 1983;
Lakshmanan et al., 2002; Bazemore et al., 2003). The key
enzymes responsible for these reactions are arginase,
adenosine deaminase, and AMP deaminase (Yeh et al.,
1978; Ward et al., 1979; Bazemore et al., 2003). The most
commonly found spoilage bacterial genera include Achro-
mobacter, Acinetobacter, Aeromonas, Alcaligenes, Bacillus,
Brevudimonas, Carnobacterium, Chryseomonas, Corynebac-
terium, Flavobacterium, Micrococcus, Moraxella; Proteus,
Pseudomonas, Serratia, Shewanella, Staphylococcus, and
Vibrio (Alvarez and Koburger, 1981; Lakshmanan et al.,
2002; Bazemore et al., 2003; Mejlholm et al., 2005).
Sensory evaluation is one of the most important items to
consider when developing a novel product or making
changes to an existing product. The product may fulll all
objectives, but if it is not accepted by the consumer, it is of
little value. Sensory analysis of shrimp products has been
conducted by several researchers (Alvarez and Koburger,
1979; Edmunds and Lillard, 1979; Matches and Layrisse,
1985; Fatima et al., 1988; Venugopal, 1993; Benner et al.,
1994; Gundavarapu et al., 1998; Bak et al., 1999; Lin et al.,
1999). These analyses usually focus on attributes most
easily noticed by the consumer such as aroma, color,
texture, and avor.
A treatment system that will reduce the growth of
spoilage bacteria and decrease or eliminate the growth of
L. monocytogenes needs to be developed for RTE shrimp.
An application of this nature should enhance safety as well
as extend shelife. This study included two objectives: (1)
to compare the combined inuence of packaging atmo-
sphere (air, vacuum, and CO2) and storage temperature (3,
7, and 12 1C) to control the growth of L. monocytogenes
and psychrotrophic spoilage bacteria on RTE shrimp; and
(2) to evaluate the sensory quality of RTE shrimp stored
under the conditions stated in objective 1.
2. Materials and methods
2.1. Inoculation of shrimp
Five strains (ATCC 15313, 51414, 43256, 19115, and
7644) of L. monocytogenes were used to make a cocktail for
inoculation. Prior to each experiment, frozen stock cultures
of each strain were individually inoculated into 200 ml of
trypticase soy broth with 0.6% yeast extract (BD, Sparks,
Maryland, USA). Once inoculated, cultures were incubated
at 32 1C for 24 h to achieve an initial starting culture of
approximately 108 cfu/ml. Two milliliters of each strain
culture were added to a sterile test tube and vortexed for
10 s to ensure thorough mixing of strains, producing 10 ml
of a ve strain cocktail. Two milliliters of cocktail were
then distributed into 2 l of 0.01 M phosphate buffered
saline (PBS, Sigma, St. Louis, Missouri, USA) in a 3 l
steamer bucket (Progressive International, Kent, Washing-
ton, USA). This distribution produced an immersion
solution containing 106 cfu/ml of L. monocytogenes.
Several 454 g packages of individually quick frozen
cooked, peeled, and deveined shrimp (100150 count) were
obtained from a local retail store and kept frozen until
used. Prior to the beginning of each experiment, packages
were placed in a 4 1C refrigerator to thaw overnight. Before
immersion into control (2 l PBS) or inoculation buffers,
four packages of shrimp were aseptically opened under a
laminar ow hood by thoroughly wiping with 70% ethanol
 ARTICLE IN PRESS
T.J. Rutherford et al. / Food Microbiology 24 (2007) 703710
and cutting horizontally below the dorsal seal with sterile
scissors. Under the laminar ow hood, 908 g of shrimp
were placed into each of two 3 l steamer baskets
(Progressive International). One basket was then dipped
into the control bucket (2 l PBS) and the other basket was
dipped into the inoculation bucket. The steamer baskets
were then immersed for 5 min with constant hand agitation
and then removed and placed on sterile paper towels to
drain for 10 min in the laminar ow hood.
2.2. Packaging
After draining, Cryovac (Duncan, South Carolina, USA)
B2650 bags (20.3 cm tall  16.5 cm wide; water vapor
transmission of 0.50.6 g per 254 cm2 at 37.8 1C, 100% relative
humidity, 24 h; oxygen transmission of 36 cc per m2 at 4.4 1C,
24 h) were aseptically lled with 20 g of shrimp and sealed
using a Multivac model A300/16 (Kansas City, Missouri,
USA). Three packaging atmospheres were used: air, vacuum,
and 100% CO2 (Nexair, Columbus, Mississippi, USA). Air
packaging was done at 999 mbar vacuum for 1 s with a 2.5 s
seal. Vacuum packaging was done at 15 mbar vacuum for 1 s
with a 2.5 s seal. Modied atmosphere packaging was done at
15 mbar vacuum for 1 s followed by a 700 mbar 100% CO2
ush and a 2.5 s seal. Packages with each atmosphere were
randomly placed into 3, 7, and 12 1C incubators for storage.
2.3. Bacterial enumeration
For each treatment, duplicate packaged shrimp samples
were analyzed for microbial counts on days 0, 3, 6, 9, 12, and
15. Twenty milliliters of PBS were added to each 20 g shrimp
sample in a stomacher bag. The bags were then homogenized
for 30 s using a stomacher (Tekmar, STO-400, Cincinnati,
Ohio, USA). Serial dilutions of the homogenate were made
in PBS and plated onto Modied Oxford Agar (MOX, BD)
and Plate Count Agar (PCA, BD) with the aid of a model D
spiral plater (Spiral Biotech, Norwood, Massachusetts,
USA). MOX plates were incubated at 28 1C for 48 h to
allow for growth of L. monocytogenes colonies, which appear
grayish-black. PCA plates were incubated at 7 1C for 5 days
to allow for growth of psychrotrophic bacteria. Microbial
counts were determined using procedures outlined in the
model D spiral plater manual.
Data consisting of log10 cfu/g of L. monocytogenes and
psychrotrophic bacteria from three replications were
analyzed using the mixed procedure of the SAS statistical
package, version 8.0 (SAS Institute Inc., Cary, North
Carolina, USA). A completely randomized design with a 3-
way factorial structure was used. This procedure allowed
for the comparison of all factors (temperature, atmosphere,
and day) utilized in this study.
2.4. Sensory evaluation
Uninoculated shrimp were packaged and stored as
previously described except that 50 g of shrimp were used
705
per package. On the day of sampling, approximately 68 g
of shrimp were placed into each of two separate
autoclaved, cleaned, and deodorized 125 ml Teon snifng
bottles (Fisher Co., Pittsburgh, Pennsylvania, USA) for
odor analysis. Random 3 digit codes were assigned to each
bottle. The two identical sets of sample bottles were each
allowed to sit at room temperature for one hour prior to
analysis. For appearance evaluation, the remaining con-
tents of each package were placed into a clear 0.95 l plastic
bag (Ziploc, Racine, Wisconsin, USA) and sealed.
A random 3 digit code was assigned to each bag. The
number codes were different on both the snifng bottles
and the bags so panelists could not correlate the two.
Sensory analysis of shrimp samples was conducted using
an 11 member panel of university employees and students.
The panelists were trained 1 h per day for 1 week prior to
data collection. During training, the panelists were given
the same number of samples that they would receive while
actually participating on the panel. Shrimp samples were
prepared in the same manner as previously described
above. After sample evaluation, a daily discussion was held
about ratings, with the package identities of the shrimp
being revealed. The panelists rated shrimp samples on a 15
point hedonic scale (15 cm) for both odor and appearance.
During training, the panelists proposed that there should
be three categories of scoring for both odor and
appearance. Scores less than 7 were termed unspoiled
and acceptable. Scores falling between 7 and 8 were
termed borderline, and scores above 8 were termed spoiled
and unacceptable.
On each panel day (30 total), panelists received nine
different samples. The nine samples consisted of air,
vacuum, and modied atmosphere packages stored at 3,
7 and 12 1C for the day being evaluated (0, 3, 6, 9, 12, or
15). For day 0, panelists were given nine differently labeled
samples that had no treatment applied. The gathering of
sensory data from ve replications was accomplished by
assigning a number to each mark placed on the line scale by
panelists for each sample. A ruler was used to measure each
mark. Thus, it was possible to calculate scores from 0 to 15.
Data were analyzed using the mixed procedure (SAS
version 8.0) to calculate least-square means for odor
and appearance scores. This procedure allowed for the
determination of product shelife as rated by the panelists
for each package, temperature, and day. A comple-
tely randomized design with a 3-way factorial structure
was used.
3. Results and discussion
3.1. Growth of L. monocytogenes
Fig. 1 illustrates the effects of different packaging
atmospheres on the growth of L. monocytogenes in RTE
shrimp stored at 3, 7, and 12 1C. Uninoculated control
samples showed no L. monocytogenes growth after 15 days
of storage at any temperature, which indicated that
 ARTICLE IN PRESS
706 T.J. Rutherford et al. / Food Microbiology 24 (2007) 703710

Air Vac Map

10
9
8
7
6
5
4
12C
3
2

10
9
7C
8
7
Log10 cfu/g

6
5
4
3
2

10
9
8 3C

7
6
5

4
3
2
0 3 6 9 12 15
TIME (days)

Fig. 1. Effect of packaging atmosphere (air, vacuum, CO2 modied atmosphere) on the growth of Listeria monocytogenes on ready-to-eat shrimp stored at
3, 7, and 12 1C. Standard error 0.02.

L. monocytogenes was not likely present in the shrimp used.
There was no signicant temperature plus atmosphere
interaction (p40.05). Thus, temperature and packaging
atmosphere acted independently on the growth of
L. monocytogenes for all three storage temperatures and
all three packaging atmospheres. Regardless of tempera-
ture, CO2 packaging was most effective (pp0.05) at
controlling the growth of L. monocytogenes, while vacuum
packaging was second most effective, and air packaging
was least effective. Overall, the most effective packaging
and temperature combination to control the growth of
L. monocytogenes was modied atmosphere combined with
storage at 3 1C.
It is widely accepted that CO2 packaging is an effective
control measure to slow the rate of L. monocytogenes
growth compared to air or vacuum packaging; however, it
is also known that the effectiveness of this treatment
declines as storage temperature increases (Marshall et al.,
1991; Hudson et al., 1994; Pothuri et al., 1995). The
importance of strict temperature control is further empha-
sized by the U.S. Food and Drug Administration, which
recommends that seafood products packaged under
vacuum or in CO2 atmospheres must be stored at
temperatures o3.3 1C to prevent growth of nonproteolytic
C. botulinum (FDA, 2001). This requirement coupled with
present data on L. monocytogenes indicates that RTE
shrimp stored under modied atmospheres should be
stored at o3.3 1C to minimize pathogen growth. If one
assumes that L. monocytogenes most likely would be a
post-processing contaminant on RTE shrimp and be
present in low numbers, using CO2 packaging and storage
at 3 1C may allow processors the opportunity to offer a
non-frozen alternative that is reasonably safe throughout
its shelife. However, additional hurdles (Marshall, 2004;
 ARTICLE IN PRESS
T.J. Rutherford et al. / Food Microbiology 24 (2007) 703710 707

Mejlholm et al., 2005) may be desirable if temperature
control is difcult during transportation, retail display, and
home use.
3.2. Growth of psychrotrophs
Fig. 2 demonstrates the effect of different packaging
atmospheres on the growth of naturally present psychro-
trophic bacteria on RTE shrimp stored at 3, 7, and 12 1C.
There was a signicant (pp0.05) temperature plus gas
interaction. Thus, growth of psychrotrophs was dependent
upon combinations of temperature and packaging atmo-
sphere. Packaging in CO2 was signicantly (pp0.05) more
effective against psychrotrophs when compared to vacuum
and air packaging.
As with L. monocytogenes, the combination of CO2
packaging with storage at 3 1C was the most effective
combination for controlling the growth of psychrotrophic
bacteria in RTE shrimp. The effect of decreased storage
temperature and CO2 atmosphere on controlling the
growth of spoilage and pathogenic organisms has been
well documented (Genigeorgis, 1985; Farber, 1991a;
Fernandez et al., 1997). The increased activity of CO2 at
low temperature is presumably due to its greater aqueous
and lipid solubility as temperature decreases (Genigeorgis,
1985; Daniels et al., 1985). When applied to biological
tissue, CO2 exists as a dissolved gas and as carbonic acid
(2%) (Daniels et al., 1985; Farber, 1991a). These two states
and the resultant decrease in pH can alter bacterial cell
membrane function to affect nutrient uptake and absorp-
tion, and can inhibit intracellular enzyme activity.
3.3. Sensory shelflife
Average sensory scores for overall odor are presented in
Fig. 3. RTE shrimp stored at 12 1C in air became
borderline by day 6 and were spoiled by day 9. When
using vacuum packaging at 12 1C, the shrimp remained

Air Vac Map

10
9
8
7
6 12C

5
4
3
2

10
9
8
Log10 cfu/g

7
6
7C
5
4
3
2

10
9
8
7
6
5 3C
4
3
2
0 3 6 9 12 15
TIME (days)

Fig. 2. Effect of packaging atmosphere (air, vacuum, CO2 modied atmosphere) on the growth of naturally present psychrotrophic bacteria on ready-to-
eat shrimp stored at 3, 7, and 12 1C. Standard error 0.05.
 ARTICLE IN PRESS
708 T.J. Rutherford et al. / Food Microbiology 24 (2007) 703710

Air Vac Map

15
14
13
12 12C
11
10 Spoiled
9
8 Borderline
7
6 Unspoiled
5
4
3
2
1
0

15
14
13
12 7C
11
10 Spoiled
9
Score

8
Borderline
7
6
Unspoiled
5
4
3
2
1
0

15
14
13 3C
12
11
10 Spoiled
9
8 Borderline
7
6 Unspoiled
5
4
3
2
1
0
0 3 6 9 12 15
TIME (days)

Fig. 3. Average odor scores of ready-to-eat shrimp stored at 3, 7, and 12 1C under different atmospheres (air, vacuum, CO2 modied atmosphere).
Standard error 0.37.

unspoiled by day 6, but were spoiled by day 9. In contrast,
no packages stored under CO2 at 12 1C reached the
borderline stage during the entire 15 day study. Likewise,
none of the shrimp stored at 3 or 7 1C in any atmosphere
were considered spoiled by day 15.
Average sensory scores for overall appearance are
presented in Fig. 4. No product became visually unac-
ceptable throughout the 15 days of storage at any
temperature or packaging atmosphere. Nevertheless, air
packs combined with a 12 1C storage temperature resulted
in borderline product on days 9 and 12. Product on the
verge of unacceptability was produced by vacuum packa-
ging and storage at 12 1C on day 12.
These sensory results were predictable. Increasing
storage temperature results in faster bacterial growth,
which in turn results in greater protein degradation and
formation of off-odors, especially malodorous amine
compounds (Lakshmanan et al., 2002; Bazemore et al.,
2003). Over extended storage time, the breakdown of
structural protein in shrimp tissue may also cause loss of
physical integrity, resulting in a mushier product with
increased purge. This appearance defect usually does
not precede off-odor formation, as was seen in the pre-
sent study.
4. Conclusions
The novelty of the present work was the focus
on refrigerated RTE shrimp. Overall, CO2 packaging
and storage at 3 1C was the most effective packaging
 ARTICLE IN PRESS
T.J. Rutherford et al. / Food Microbiology 24 (2007) 703710 709

Air Vac Map

15
14
13 12C
12
11
10
9 Unacceptable
8
7 Borderline
6
5 Acceptable
4
3
2
1
0

15
14
13 7C
12
11
10 Unacceptable
9
Score

8 Borderline
7
6 Acceptable
5
4
3
2
1
0

15
14
13 3C
12
11
10 Unacceptable
9
8 Borderline
7
6 Acceptable
5
4
3
2
1
0
0 3 6 9 12 15
TIME (days)

Fig. 4. Average appearance scores of ready-to-eat shrimp stored at 3, 7, and 12 1C under different atmospheres (air, vacuum, CO2 modied atmosphere).
Standard error 0.44.

system for controlling growth of both L. monocytogenes Acknowledgments

and other psychrotrophic bacteria as well as offering
an extended shelife compared to air and vacuum
packaging. This packaging combination also would con-
trol the growth of another psychrotrophic pathogen,
nonproteolytic C. botulinum. Storage temperatures of
greater than 3.3 1C would not control either pathogen.
For practical reasons, strict temperature control is
often difcult to achieve during the entire course of
processing, warehousing, shipping, retail display, and
home storage. The present study demonstrates that
even small increases in storage temperature can have a
dramatic effect on the growth of L. monocytogenes
while not impacting shelife when stored under CO2.
Further research should be conducted with this pro-
duct to nd additional antimicrobial barriers that may
provide added protection during temperature abuse
scenarios.
Approved for publication as journal article no. J-11065
of the Mississippi Agricultural and Forestry Experiment
Station. This work was supported in part by USDA-
CSREES special Grant no. 2003-34231-13064, by the
Mississippi Agricultural and Forestry Experiment Station
under project MIS-081430, and by the National Fisheries
Institute.
References
Alvarez, R.J., Koburger, J.A., 1979. Effect of delayed heading on some
quality attributes of Penaeus shrimp. J. Food Prot. 42, 407409.
Alvarez, R.J., Koburger, J.A., 1981. Effect of Planococcus citreus on
selected quality indices of Penaeus shrimp. J. Food Prot. 44, 359363.
Bak, L.S., Andersen, A.B., Andersen, E.M., Bertelsen, G., 1999. Effect of
modied atmosphere packaging on oxidative changes in frozen stored
cold water shrimp (Pandalus borealis). Food Chem. 64, 169175.
 ARTICLE IN PRESS
710 T.J. Rutherford et al. / Food Microbiology 24 (2007) 703710
Bazemore, R., Fu, S.G., Yoon, Y., Marshall, D., 2003. Major causes of
shrimp spoilage and methods for assessment. In: Rimando, A.M.,
Schrader, K.K. (Eds.), Off-Flavors in Aquaculture. ACS Symposium
Series No. 848. American Chemical Society, Washington, DC, USA,
pp. 223234.
Benner, R.A., Miget, R., Finne, G., Acuff, G.R., 1994. Lactic acid/
melanosis inhibitors to improve shelf life of brown shrimp (Penaeus
aztecus). J. Food Sci. 59, 242245, 250.
Buchanan, R.L., 1991. Microbiological criteria for cooked, ready-to-eat
shrimp and crabmeat. Food Technol. 45 (4), 157160.
Chang, O., Cheuk, W.L., Nickelson, R., Martin, R., Finne, G., 1983.
Indole in shrimp: effect of fresh storage temperature, freezing and
boiling. J. Food Sci. 48, 813816.
Daniels, J.A., Krishnamurthi, R., Rizvi, S.S.H., 1985. A review of effects
of carbon dioxide on microbial growth and food quality. J. Food Prot.
48, 532537.
Dillon, R.M., Patel, T.R., 1992. Listeria in seafoods: a review. J. Food
Prot. 55, 10091015.
Dorsa, W.J., Marshall, D.L., 1995. Inuence of lactic acid and modied
atmosphere on thermal destruction of Listeria monocytogenes in
crawsh tail meat homogenate. J. Food Saf. 15, 19.
Dorsa, W.J., Marshall, D.L., Moody, M.W., Hackney, C.R., 1993. Low
temperature growth and thermal inactivation of Listeria monocyto-
genes in precooked crawsh tail meat. J. Food Prot. 56, 106109.
Duran, G.M., Marshall, D.L., 2005. Ready to eat shrimp as an
international vehicle of antibiotic resistant bacteria. J. Food Prot.
68, 23952401.
Edmunds, W.J., Lillard, D.A., 1979. Sensory characteristics of oysters,
clams, and cultured and wild shrimp. J. Food Sci. 44, 368373.
Farber, J.M., 1991a. Microbiological aspects of modied-atmosphere
packaging technologya review. J. Food Prot. 54, 5870.
Farber, J.M., 1991b. Listeria monocytogenes in sh products. J. Food
Prot. 54, 922924.
Fatima, R., Khan, M.A., Qadri, R.B., 1988. Shelf life of shrimp (Penaeus
merguiensis) stored in ice (0 1C) and partially frozen (3 1C). J. Sci.
Food Agric. 42, 235247.
FDA, 2001. Fish and sheries products hazards and controls guidance:
third edition. U.S. Food and Drug Administration, CFSAN, Ofce of
Seafood, Rockville, Maryland, USA.
Fernandez, P.S., George, S.M., Sills, C.C., Peck, M.W., 1997. Predictive
model of the effect of CO2, pH, temperature, and NaCl on the growth
of Listeria monocytogenes. Int. J. Food Microbiol. 37, 3745.
Gecan, J.S., Bandler, R., Staruskiewicz, W.F., 1994. Fresh and frozen
shrimp: a prole of lth, microbiological contamination, and decom-
position. J. Food Prot. 57, 154158.
Genigeorgis, C.A., 1985. Microbial and safety implications of the use of
modied atmospheres to extend the storage life of fresh meat and sh.
Int. J. Food Microbiol. 1, 237251.
Gimenez, B., Roncales, P., Beltran, J.A., 2002. Modied atmo-
sphere packaging of lleted rainbow trout. J. Sci. Food Agric. 82,
11541159.
Gundavarapu, S., Hung, Y.C., Reynolds, E.A., 1998. Consumer
acceptance and quality of microwave-cooked shrimp. J. Food Qual.
21, 7184.
Hathed, A.A.M., Maqbool, T.K., Kumar, S.S., 2003. Microbial quality of
shrimp products of export trade produced from aquacultured shrimp.
Int. J. Food Microbiol. 82, 213221.
Hudson, J.A., Mott, S.J., Penney, N., 1994. Growth of Listeria
monocytogenes, Aeromonas hydrophila, and Yersinia enterocolitica on
vacuum and saturated carbon dioxide controlled atmosphere-pack-
aged sliced roast beef. J. Food Prot. 57, 204208.
Lakshmanan, R., Shakila, R.J., Jeyasekaran, G., 2002. Survival of amine-
forming bacteria during the ice storage of sh and shrimp. Food
Microbiol. 19, 617625.
Lannelongue, M., Finne, G., Hanna, M.O., Nickelson, R., Vanderzant,
C., 1982. Storage characteristics of Brown shrimp (Penaeus aztecus)
stored in retail packages containing CO2-enriched atmospheres. J.
Food Sci. 47, 911914.
Layrisse, M.E., Matches, J.R., 1984. Microbiological and chemical
changes of Spotted shrimp (Pandalus platyceros) stored under modied
atmospheres. J. Food Prot. 47, 453457.
Lin, T., Durance, T.D., Scaman, C.H., 1999. Physical and sensory
properties of vacuum microwave dehydrated shrimp. J. Aquat. Food
Prod. Technol. 8 (4), 4153.
Lionberg, W., Restaino, L., Frampton, E.W., Lewis, V., Liedtke, M.,
2003. Listeria monocytogenes incidence and distribution on a selective/
differential plating medium in Listeria-positive environmental samples
from ready-to-eat food facilities. Food Prot. Trends 23, 554557.
Marshall, D.L., 2004. Control of Listeria monocytogenes in ready-to-eat
foods. In: Beier, R.C., Pillai, S.D., Phillips, T.D., Ziprin, R.L. (Eds.),
Pre-Harvest and Post-Harvest Food Safety: Contemporary Issues and
Future Directions. Institute of Food Technologists and Blackwell
Publishing, Ames, Iowa, USA, pp. 349364.
Marshall, D.L., Wiese-Lehigh, P.L., Wells, J.H., Farr, A.J., 1991.
Comparative growth of Listeria monocytogenes and Pseudomonas
fluorescens on precooked chicken nuggets stored under modied
atmospheres. J. Food Prot. 54, 841843, 851.
Marshall, D.L., Andrews, L.S., Wells, J.H., Farr, A.J., 1992. Inuence of
modied atmosphere packaging on the competitive growth of Listeria
monocytogenes and Pseudomonas fluorescens on precooked chicken.
Food Microbiol. 9, 303309.
Matches, J.R., Layrisse, M.E., 1985. Controlled atmosphere storage of
Spotted shrimp (Pandalus platyceros). J. Food Prot. 48, 709711.
Mejlholm, O., Bkns, N., Dalgaard, P., 2005. Shelf-life and safety
aspects of chilled cooked and peeled shrimps (Pandalus borealis) in
modied atmosphere packaging. J. Appl. Microbiol. 99, 6676.
Motes, M.L., 1991. Incidence of Listeria spp. in shrimp, oysters, and
estuarine waters. J. Food Prot. 54, 170173.
NOAA, 2003. Fisheries of the United States: 2002. U.S. Department of
Commerce, National Marine Fisheries Service, Silver Springs, Mary-
land, USA.
Notermans, S., Hoornstra, E., 2000. Risk assessment of Listeria
monocytogenes in sh products: some general principles, mechanism
of infection and the use of performance standards to control human
exposure. Int. J. Food Microbiol. 62, 223229.
Oh, D.H., Marshall, D.L., 1995. Inuence of packaging method, lactic
acid and monolaurin on Listeria monocytogenes in crawsh tail meat
homogenate. Food Microbiol. 12, 159163.
Pothuri, P., Marshall, D.L., McMillin, K.W., 1995. Combined effects of
packaging atmosphere and lactic acid on growth and survival of
Listeria monocytogenes in craysh tail meat at 4 1C. J. Food Prot. 59,
253256.
Shineman, T.L., Harrison, M.A., 1994. Growth of Listeria monocytogenes
on different muscle tissues. J. Food Prot. 57, 10571062.
Suwansonthichai, S., Rengpipat, S., 2003. Enumeration of coliforms and
Escherichia coli in frozen black tiger shrimp Penaeus monodon by
conventional and rapid methods. Int. J. Food Microbiol. 81, 113121.
Valdimarsson, G., Einarsson, H., Gudbjornsdottir, B., Magnusson, H.,
1998. Microbiological quality of Icelandic cooked-peeled shrimp
(Pandalus borealis). Int. J. Food Microbiol. 45, 157161.
Venugopal, V., 1993. Cook-chill process to extend refrigerated shelf life of
peeled and deveined shrimp and white pomfret. Int. J. Food Sci.
Technol. 28, 273278.
Ward, D.R., Finne, G., Nickelson II, R., 1979. Use of a specic-ion
electrode (ammonia) in determining the quality of shrimp. J. Food Sci.
44, 10521054, 1057.
Yeh, C.S., Nickelson II, R., Finne, G., 1978. Ammonia-producing
enzymes in white shrimp tails. J. Food Sci. 43, 14001401, 1404.