Oxygen depletion results from the combination
of several physical and biological processes. In the Gulf of Mexico, hypoxia
results from the stratification of marine waters due to Mississippi River
system freshwater inflow and the decomposition of organic matter stimulated
by Mississippi River nutrients. As a general rule, the nutrients delivered
to estuarine and coastal systems support biological productivity. Excessive
levels of nutrients, however, can cause intense biological productivity
that depletes oxygen. The remains of algal blooms and zooplankton fecal
pellets sink to the lower water column and seabed. The rate of depletion
of oxygen during processes that decompose the fluxed organic matter exceeds
the rate of production and resupply from the surface waters, especially
when waters are stratified. Stratification in the northern Gulf of Mexico
is most influenced by salinity differences year-round, but is accentuated
in the summer due to solar warming of surface waters and calming winds.
Following a fairly predictable annual cycle beginning in the spring, oxygen
depletion becomes most widespread, persistent and severe during the summer
months.
Midsummer coastal hypoxia in the northern
Gulf of Mexico was first recorded in the early 1970s. In recent years (1993-1999),
the extent of bottom-water hypoxia (16,000 to 20,000 km2, see Figure 1) has
been greater than twice the surface area of the Chesapeake Bay, rivaling
extensive hypoxic/anoxic regions of the Baltic and Black Seas. Prior to
1993, the hypoxic zone averaged 8,000 to 9,000 km2 (1985-1992)
(Rabalais et al., 1998, 1999). The hypoxic area covered 12,400 km2
in 1998, about the size of Connecticut. Maps of the extent of bottom-water midsummer
hypoxia
(see examples in Figure 2)
provide a benchmark for year-to-year comparisons, but should not
be overinterpreted (Rabalais et al., 1999). The difference in hypoxia
in a drought (1988) versus a flood (1993) year suggests a relationship
between river discharge and the extent of hypoxia. A simple linear regression
of midsummer area and mean Mississippi River discharge for the preceding
year produced an R2 of 0.934 for
nine years of data from 1985-1993 (Wiseman et al., 1997). This relationship,
however, fails to hold for the additional years of 1994-1998. Thus, a comparison
of midsummer area (a minimal and rough estimate) versus discharge is not
entirely satisfactory. There is evidence, for example, that carbon burial
in 1993 was sufficient to support the extensive 1994 hypoxic zone despite
"normal" flow conditions (Justic et al.,
1997). Examples of variability in the midsummer extent of hypoxia due to
physical conditions at the time of sampling include the reduced midsummer
size in 1997 and 1998 due to the passage of a hurricane on the southeastern
portion of the study area, and the current regime, respectively. Stronger
relationships are evident for hypoxia versus river discharge and nutrient
flux given time and spatial lags.
The persistence of the extensive midsummer
areas is known from a few back-to-back shelfwide cruises in 1993 and 1994 (see Figure 3).
These cruises found that the large size of the hypoxic zone persisted over 2- to 3-week
periods, but varied somewhat in configuration. Hypoxia occurs from late February through early
October, nearly continuously from mid-May through mid-September, and is
most widespread, persistent, and severe in June, July, and August. Hypoxic
waters can include 20 to 80% of the lower water profile between 5 and 30 m water
depth, and can extend as far as 130 km offshore. Throughout its distribution,
the impact of hypoxic bottom waters is exacerbated by the release of toxic
hydrogen sulfide from sediments (Harper et al., 1981, 1991).
The timing and location of low dissolved oxygen
conditions in coastal waters is now fairly well documented, and there are
studies that link the frequency and volume of summer oxygen depletion to
increased nutrient inputs (Officer et al., 1984; Larsson et al., 1985;
Tolmazin, 1985; Andersson and Rydberg, 1987; Justic et al., 1987; Cooper
and Brush, 1991, 1993; Diaz and Rosenberg, 1995; Rabalais
et al., 1996). Growth in population, changes in land cover, increase in
agricultural acreage, and increases in fertilizer use and animal husbandry
have resulted in two- to tenfold increases in the level of nutrient inputs
during this century, with particularly dramatic increases since the 1950s
(Turner and Rabalais, 1991; Justic et al., 1995a, 1995b; Howarth et al., 1996; Nixon, 1997, Goolsby et al., 1999).
Studies in the northern Gulf of Mexico, among
other systems, provide evidence that the timing, spatial extent, and severity
of oxygen depletion, as well as changes over time, are linked to freshwater
discharge and nutrient flux (Justic et al., 1993, 1996, 1997; Rabalais et al.,
1996). The severity and duration
of oxygen depletion in the northern Gulf of Mexico depend, at least partially,
on the amplitude and phasing of discharge from the Mississippi and Atchafalaya
Rivers (Justic et al., 1993; Rabalais
et al., 1996, 1998, 1999). Figure 3 illustrates the high variability of hypoxia possible
during a one month time frame. The timing of Mississippi River discharge and
nutrient flux, surface water phytoplankton production, and oxygen depletion
are strongly correlated on the southeastern Louisiana shelf (Lohrenz et
al., 1997, Justic et al., 1997).
The highest surface water productivity occurs one month after the maximum
river discharge, and oxygen depletion in bottom waters occurs two months
after the highest river discharge (Justic et al., 1993).
Figure 4 illustrates the increased frequency of hypoxia down current from the discharges of the Mississippi and Atchafalaya Rivers.
Mississippi River nutrient concentrations and
loadings to the adjacent continental shelf changed dramatically during
this century, with an acceleration of these changes since the 1950s (Turner
and Rabalais, 1991; Justic et al.,
1995a, 1995b). Nitrogen is the principal nutrient yielding excess organic
matter sedimentation to the Gulf hypoxic zone. Nitrogen export from the
Mississippi River system has increased two- to sevenfold over the last century
(Turner and Rabalais, 1991; Goolsby et al., 1999; Howarth et al., 1996).
The majority of Mississippi River nitrogen originates from agricultural
practice, while smaller fractions arise from human sewage, nonagricultural
fertilizer use, and precipitation (Howarth et al., 1996; Goolsby et al.,
1999; Downing et al., 1999). Silica and phosphorus also play a role, and
the changing balance of nitrogen, silica, and phosphorus can affect marine
food webs (Turner et al., 1998).
Hypoxia was first recorded on the continental
shelf of the northern Gulf of Mexico in the early 1970s. A few directed
studies in 1975-1976 (Ragan et al., 1978; Turner and Allen, 1982) and a
series of environmental assessments revealed midsummer hypoxia in many
inner shelf areas. Consistent data collection on the distribution and dynamics
of hypoxia began in 1985 (Rabalais et al., 1991). Prior to the 1970s, scattered
anecdotal data from shrimp trawlers in the 1950s to 1960s reported low
or no catches, which was considered consistent with hypoxia but may also
reflect other causes. Analysis of the sediment record, however, shows that
severe hypoxia is a recent development in Gulf waters.
The sedimentary record supports the inference of increased eutrophication
(increased rate of primary production) and hypoxia in the Mississippi River delta
bight, primarily because of changes in nitrogen loading (Turner and Rabalais,
1994; Eadie et al., 1994; Nelsen et al., 1994; Sen Gupta et al., 1996;
Rabalais et al., 1996). Changes in the productivity of surface waters and
oxygen deficiency over time can be reconstructed from chemical and biological
analyses of bottom sediments. In several U.S. estuaries and coastal waters,
there is evidence of decade- and century-long increases in nutrient enrichment,
hypoxia, and eutrophication. In the northern Gulf of Mexico, for example,
sediment analysis shows clearly that the productivity of surface waters
adjacent to the Mississippi River has risen as the nitrogen level in the
river has risen (Eadie et al., 1994; Turner and Rabalais, 1994). Higher
surface water production results in larger carbon fluxes to the bottom,
thus increasing the severity and spatial extent of hypoxia (Qureshi, 1995;
Rabalais et al., 1998). Analysis of benthic organisms, foraminifera, indicates
that oxygen stress has worsened this century and accelerated since the
1950s when nitrogen flux in the river began to increase (Nelsen et al.,
1994; Rabalais et al., 1996; Sen Gupta et al., 1996). These results are
consistent with similar long-term evidence of eutrophication and worsening
oxygen stress worldwide. The problems the Gulf of Mexico faces from hypoxia
are not unique. Hypoxia related to human activities currently threatens
many of the major coastal embayments and estuaries in the world (Diaz and
Rosenberg, 1995).
Hypoxia affects living resources, biological
diversity, and the capacity of aquatic systems to support biological populations.
When oxygen levels fall below critical values (2 mg/l, for the northern
Gulf of Mexico), those organisms capable of swimming (e.g., demersal fish,
portunid crabs, and shrimp) evacuate the area (see Figure 5). The stress on less motile
fauna caused by declining oxygen levels varies according to the oxygen
requirements of the organism. Animals resident in the seabed are usually
more resistant to low oxygen concentrations, but they also experience stress
or die as oxygen concentrations decline from 1 mg/l to anoxia. Important
fishery resources are variably affected by direct mortality, forced migration,
reduction in suitable habitat, increased susceptibility to predation, changes
in food resources and disruption of life cycles. Commercial and recreational
fisheries in the Gulf generate $2.8 billion annually (Holiday and O'Bannon,
1997). Signs consistent with impact on Gulf of Mexico fisheries are
(1)reduced food resources for fish and shrimp in hypoxic waters (Rabalais
et al., 1995);
(2) reduced abundance of fish and shrimp in hypoxic waters
(Pavela et al., 1983; Leming and Stuntz, 1994; Renaud, 1986);
(3) declinein shrimp catch and catch efficiency since hypoxia expanded (Zimmerman
et al., 1997; Downing et al., 1999); and
(4) loss of production potential due
to the blocked migration of juvenile shrimp offshore by the presence of
the hypoxic zone (Nance et al., 1994; Zimmerman et al., 1997; Downing et
al., 1999).
The northern Gulf of Mexico adjacent to the discharge of the Mississippi
River system is an example of a coastal ocean that has undergone eutrophication
as a result of increasing nutrients and that has worsened century-long and accelerating
recent decadal hypoxic conditions. Models that link Mississippi River discharge
with Gulf of Mexico hypoxia demonstrate worsening hypoxia in bottom waters
with increased freshwater discharge and even worse hypoxia with additional
nitrogen accompanying the increased discharge (Bierman et al., 1994;
Justic et al., 1996, 1997).
Conversely, the models show that a reduction in oxygen demand
in the lower water column will result from a reduction in nitrogen (and
to a lesser degree the phosphorus) load to the surface waters (Limno-Tech,
Inc., 1995). In other words, hypoxia in the northern Gulf of Mexico can
be alleviated to some degree by a reduction in the nutrient loading. Whole
system management of the entire watershed where most of the changes have
occurred over the last several decades is a necessary step in alleviating
the problems in the Gulf of Mexico. The ability to detect changes in the
coastal system (given any nutrient reductions) will be complicated by an
inherently variable biological system and extreme events. In addition,
the eutrophic state may be persistent and recovery may be slow. Still,
there are several success stories for improvement of estuarine and coastal
ecosystems in response to nutrient abatement in the watershed or in direct
discharges to the system, and similar activities on larger coastal systems
with a much larger watershed, while daunting, are worthwhile and achievable.
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Nancy N. Rabalais is a professor at the Louisiana Universities Marine Consortium,
where she has been working since 1983. She earned a Ph.D. in zoology from The
University of Texas at Austin in 1983, and her B.S. and M.S. from Texas A&I University,
Kingsville, in 1972 and 1975. Before she joined LUMCON, Dr. Rabalais was a research associate,
then graduate student, at the U.T. Marine Science Institute, Port Aransas Marine Laboratory.
She teaches marine science courses at LUMCON and in the Department of Oceanography and
Coastal Sciences at Louisiana State University. Dr. Rabalais' research interests include
the dynamics of hypoxic environments, interactions of large rivers with the coastal ocean,
estuarine and coastal eutrophication, benthic ecology, and the environmental effects of
habitat alterations and contaminants. Dr. Rabalais is a Fellow of the American Association
for the Advancement of Science, an Aldo Leopold Leadership Fellow, a co-recipient of the
1999 Blasker Award for Environmental Science and Engineering, a NOAA Environmental Hero,
and a past-President of the Estuarine Research Federation, and a member of the Ocean Studies Board of the National Research Council (2000-2003).
A large zone of oxygen-depleted
water extends across the Louisiana continental shelf and on to the Texas
coast most summers. The Gulf of Mexico hypoxic zone is the largest such
zone in coastal waters of the Western Hemisphere. The occurrence of severe
oxygen depletion, either hypoxia (< 2 mg/l, or < 3 mg/l in some systems)
or anoxia (0 mg/l), is a growing concern for U.S. estuarine and coastal
waters. Many hypoxic zones elsewhere in the world have been caused by excess
nutrients exported from rivers, resulting in reduced commercial and recreational
fisheries. Prolonged oxygen depletion not only disrupts benthic and demersal
communities but can cause mass mortalities of aquatic life (Diaz and Rosenberg,
1995). Among other problems, the consequences to coastal commercial fisheries
can be disastrous (Baden et al., 1990; Zaitsev, 1991, 1993). Hypoxic zones
are now one of the most widespread and accelerating human-induced deleterious
impacts on the world's marine environments.