(Received 23 February 2010; in revised form 4 July 2010; accepted 19 August 2010)
Measurements of the specific absorption coefficients of phytoplankton (a*ph) are cur- Keywords:
rently required to estimate primary productivity at regional to global scales using Specific absorp-
satellite imagery. The variability in a*ph and phytoplankton size fraction was deter- tion,
mined during January 2002 in the southern region of the California Current. Median phytoplankton,
flow cytometry,
values of a* ph at 440 nm and 674 nm were 0.061 and 0.028 m2 (mg Chl-a)1 and sig-
HPLC-pigments,
nificant variability was found between inshore and offshore stations. A decrease of
California Current.
a* ph is associated with increased phytoplankton abundance and larger species. The
a* ph tends to be high when the photoprotector zeaxanthin is present in elevated con-
centrations and phytoplankton abundance lower. The nano-microphytoplankton (>5
m) community consisted of 28 diatom and 15 dinoflagellate genera with mean abun-
dance values of 2.8 and 1.6 103 cells l1, respectively. The picophytoplankton (<5
m) community consisted of Prochlorococcus sp. (mean 8.2 10 6 cells l 1 ) and
Synechococcus sp. (mean 19.5 106 cells l1), as well as a mixture of picoeukaryotes
(mean 8.6 10 6 cells l 1 ). The contributions of nano-microphytoplankton and
picophytoplankton to the total biomass ( g C l 1) were 46% and 54%, respectively.
This study showed that picophytoplankton cells increased 2.5 times up during Janu-
ary 2002 compared with the previous year. It was concluded that the waning of La
Nia conditions had a clear effect on the pelagic ecosystem in January 2002 and that
the higher microphytoplankton abundance in the California Current was dominated
by local and regional seasonal processes.
719
32
1 ENSENADA
2
3
4
5 14
GU
6
7 13
LF
0 12
10 11
OF
10 5
9 161
C
30 03 8 17 PUNTA
t1
AL
sec 18 BAJA
Tra
n 19 27
IFO
21 20
7 26
10
RN
25
24 28
29
IA
GUADALUPE 23
2 30 45
ISLAND 110 2 31 44
32
33 46
13 34 43
ct 1 42
28 nse 41 47
Tra 39 48
40
38 49 PUNTA
50
51
EUGENIA
37
36 52
53 62
117 35 61
54
55 60
58 59 65 63
56 64
0 57 23
12 1 66
n s ect 73
26 Tra 68 72
67
71
1 27
70
30 69
t1
n sec
Tra
U. S. A.
24
MEXICO
IMECOCAL Grid
Pacific Ocean 0 100 200 km
Fig. 1. Study area and location of stations sampled during the January 2002 cruise off Baja California. Samples were taken at 10
m depth.
attributed this variability to the relation between photo- ters. Figure 1 shows the location of the 10 transects and
synthetic and photoprotective carotenoids, blue-red ratio 73 hydrographic stations (37 km between stations and 74
of phytoplankton absorption, and sizes of phytoplankton km between transects); data taken at each station include
species. The goal of this study is to understand the vari- conductivity-temperature-depth (CTD/rosette) casts to
ability in a*ph relative to changes in the phytoplankton 1000 m water depth permitting. Samples were collected
community structure during January 2002 in the south- in coastal/oceanic waters off Baja California between 19
ern region of the California Current. January and 06 February 2002 at a depth of 10 m with 5-
litre polyethylene Niskin bottles using Tygon tubing.
2. Materials and Methods Particulate matter was filtered through Whatman GF/F
filters for the analysis of spectral absorption coefficients
2.1 Sampling and storage and phytoplankton pigment. Samples for microscope
The Mexican California Current research program analysis were stored in 250 ml dark bottles and preserved
(IMECOCAL, acronym for Investigaciones Mexicanas de with formalin-borate (0.4% final concentration). Seawater
la Corriente de California) was designed to occupy the samples of 4 ml were preserved with glutaraldehyde (0.1%
historical CalCOFI (California Cooperative Oceanic Fish- final concentration) and stored in liquid nitrogen until
eries Investigations (CalCOFI) transects in Mexican wa- analysis by flow cytometry.
2.2 Absorption coefficient of phytoplankton nm at 1-nm resolution, and repeated after rinsing the fil-
Samples for light absorption by phytoplankton were ters twice with warm methanol (Kishino et al., 1985) for
filtered onto Whatman GF/F glass microfiber filters (25 15 minutes. The absorption spectra were corrected for two
mm, 0.7 m). Sample filters were put into tissue capsules factors: baseline from blank filters and path-length am-
(Fisher HistoPrep) and then stored in liquid nitrogen for plification () by adjusting the optical density of the fil-
a maximum of one month until analysis. Blank filters were tered samples (ODfil()) to the optical density of samples
saturated with seawater through 0.2-m Nuclepore fil- in suspension (ODsus()) (Eq. (1)), which were previously
ters. Absorption of sample and blank filters was measured determined using the spectrophotometer. Phytoplankton
on a Perkin-Elmer Lambda 10 spectrophotometer with pigment absorption (aph()) was determined as the dif-
integrating sphere, following the procedure proposed by ference between total particulate matter absorption (ap())
Mitchell (1990) and Cleveland and Weidemann (1993). and non-pigmented or detritus material absorption
Sample absorption was measured between 400 and 750 (adet()) (Eq. (2)). The specific absorption coefficient of
Specific Absorption Coefficient and Phytoplankton Community in the Southern California Current 721
32
Ensenada
31
0.125
Punta
Baja 0.115
30
0.105
0.095
29
0.085
0.075
28
Punta 0.065
Eugenia 0.055
27 0.045
0.035
26 0.025
25
a* at 440nm [m (mg Chl-a ) ]
ph
2 -1
Fig. 2. Spatial distribution of the specific absorption coefficient (a*ph at 440 nm) (m2(mg Chl-a)1) at 10 m depth during January
2002.
phytoplankton (a* ph) was obtained by normalizing the sp., mean cell biovolumes of 0.52, 3.05, and 0.27 m3,
phytoplankton absorption data (m1) by the concentration respectively, were assumed.
of chlorophyll-a (Chl-a, mg m 3) measured by high-per-
formance liquid chromatography (HPLC) (Eq. (3)): 2.4 Flow cytometry
Samples were processed through a FACScan flow
OD sus = 0.368OD filt + 0.4068(ODfilt)2 (1) cytometer (Becton-Dickinson) with an air-cooled argon
(488 nm) laser at high flow rate (ca. 60 l min1) for 5
aph() = ap() a det() (2) min or until 106 events had been recorded. Data were ac-
quired by triggering on chlorophyll fluorescence (FL3)
a*ph() = aph()/Chl-a. (3) in log mode. As an internal standard, we added 10 l per
600 l sample of a 105 ml1 solution of yellow-green 0.92
2.3 Microscope observation of phytoplankton m diameter Polysciences latex beads. Synechococcus sp.
Within two months of sampling, 50 ml of water were cells were detected by the orange fluorescence signature
allowed to settle in a composite chamber and examined of phycoerythrin (FL2) versus red chlorophyll fluores-
using the inverted microscope technique (Utermhl, cence (FL3). Prochlorococcus sp. were distinguished by
1958). Diatoms and dinoflagellates (>5 m) were classi- a lower FL3 signal and no FL2 signal. Autotrophic
fied to the lowest possible taxonomic level, and identi- picoeukaryotes had higher FL3 and scatter signals and
fied based on Licea et al. (1996) and Moreno et al. (1997). low or no FL2 signal.
Only in the case of diatoms, the cells (length and width)
were measured using a micrometer. These measurements 2.5 HPLC analysis of pigments
of the nano-microphytoplankton size fractions were con- Chlorophylls and carotenoids were separated at the
verted to biovolume assuming the steriometric shapes Center for Hydro-Optics and Remote Sensing (CHORS)
suggested by Strathmann (1967) and Edler (1979). The at San Diego State University, using the Wright et al.
equation proposed by Verity et al. (1992) was then used (1991) method, which separates the major pigment com-
to convert volume to carbon concentration. For pounds found in phytoplankton. An internal pigment
Synechococcus sp., picoeukaryotes, and Prochlorococcus standard canthaxanthin was added to the 90% acetone
Stations
0.06
0.04
0.02
0.00
400 450 500 550 600 650 700 750
Wavelength (nm)
Fig. 3. Spectral magnitude of specific absorption by phytoplankton at 10 m depth during January 2002. The numbers to the right
of the arrow show the position sequence of the stations.
32
Ensenada
40000
37000
31
34000
31000
30
Punta 28000
Baja 25000
22000
19000
29
16000
13000
10000
28
7000
Punta 4000
Eugenia
1000
27
0
26
25
Nano-microphyto -1
> 5 m cells l
-118 -117 -116 -115 -114 -113 -112
Fig. 4. Spatial distribution of nano-microphytoplankton (>5 m cells l1) at 10 m depth during January 2002. Abundance esti-
mated by inverted microscopy.
sample to correct for volume changes during the extrac- resolution) and a fluorescence detector for degradation
tion process. Pigment peaks were detected by a two-chan- products. Although the absorption peaks for monovinyl
nel absorption detector (436 and 450 nm), a scanning di- Chl-a and divinyl Chl-a co-elute using this method, each
ode array absorption detector (190 to 800 nm at 1 nm compound absorbs differently at 436 and 450 nm and it
Specific Absorption Coefficient and Phytoplankton Community in the Southern California Current 723
32
Ensenada
31
200
175
30 Punta
Baja
150
29 125
100
28
750
Punta
500
Eugenia
27
250
0
26
25
Picophyto -1
< 5 m x 10 cells l
6
Fig. 5. Spatial distribution of picophytoplankton (<5 m 10 6 cells l 1) (prochlorophytes, picoeukaryotes, and cyanobacteria) at
10 m depth during January 2002. Abundance estimated by flow cytometry.
is therefore possible to correct for divinyl Chl-a contami- Thalassionema sp., and Chaetoceros spp., while the most
nation by monitoring changes in this ratio as a function common dinoflagellates were Gymnodinium spp.,
of changes in the divinyl percentages (Latasa et al., 1996). Gyrodinium spp., and Prorocentrum sp. High diatom
abundances of ~37.8 103 cells l 1 were observed at sta-
3. Results tions 59, 68, and 69. Dinoflagellate abundances ranged
from 0.2 to 25 103 cells l1 off Baja California, with
3.1 Specific absorption coefficient of phytoplankton higher values at stations 11 and 13 in the northern area,
At 10 m depth during the survey period, a*ph at 440 and at station 64 to the south.
-nm (a* ph (440)) varied between 0.024 and 0.126 m 2 The abundance of picophytoplankton (<5 m) was
(mg Chl-a)1, while a*ph at 674 nm (a*ph(674)) varied be- ~50 106 cells l1 in the study area, except at stations 1,
tween 0.008 and 0.065 (Table 1). The minimum values of 2, 47, 48, 50, 51, and 58 where values were above 50
a*ph(440) were found at station 13, located to the south 106 and up to 195 106 cells l 1 (Fig. 5). The most abun-
of Ensenada, and at stations 58, 68, and 69 off Punta dant organism was Synechococcus sp. with values rang-
Eugenia; this parameter is not homogeneous, and it ing from 0.02 to 132 10 6 cells l 1 , followed by
presents some small areas with higher and lower values picoeukaryotes with 0.2 to 60 10 6 cells l 1 , while
(Fig. 2). The magnitude of the spectral absorption curves Prochlorococcus sp. ranged from not detected to 57 106
showed greater variability in the blue band (440 nm) than cells l 1 . The contributions of the size-fractionated
in the red band (674 nm) (Fig. 3). The blue red peak ratio biomass to total phytoplankton ( g C l1) were 36% for
(aph 440 nm/a ph 674 nm) ranged from 1.70 to 4.45 (Table microphytoplankton, 32% for picoeukaryotes, 16% for
1). Synechococcus sp., 10% for nanophytoplankton, and 6%
for Prochlorococcus sp. (Fig. 6). The smaller cells
3.2 Phytoplankton community structure (picophytoplankton) made up the dominant size fraction,
Abundances of nano-microphytoplankton (>5 m) contributing 54% of the total phytoplankton biomass.
were highest at the farshore stations off Punta Eugenia,
on transects 123 and 127 (Fig. 4). Taxonomic analyses 3.3 Pigments
revealed a total of 28 diatom and 15 dinoflagellate gen- Zeaxanthin is a characteristic pigment of
era. The most common diatoms were Nitzschia spp., Synechococcus sp. while divinyl Chl-a is a unique pig-
32
Ensenada
31 0.1
0.09
Punta
30 Baja 0.08
0.07
29 0.06
0.05
28 0.04
Punta 0.03
Eugenia
0.02
27
0.01
0
26
25 Zeaxanthin (mg m -3 )
Fig. 7. Spatial distribution of zeaxanthin (mg m3) at 10 m depth during January 2002. Pigments concentration estimated by
HPLC.
Specific Absorption Coefficient and Phytoplankton Community in the Southern California Current 725
Table 2. Pigment concentrations (mg m3). Chlorophyll-a (Chl-a), zeaxanthin (Zea), divinyl chlorophyll-a (DV Chl-a), from 10
m samples collected during January 2002.
0.10
Zeaxanthin (mg m-3)
0.08
0.06
0.04
0.02 n = 64
Correlation: r = 0.76
0.00
0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 0.16 0.18 0.20
Specific Absorption Coefficient and Phytoplankton Community in the Southern California Current 727
32
Ensenada
31
19.5
19
Punta
30 18.5
Baja
18
29 17.5
17
16.5
28
16
Punta 15.5
Eugenia
27 15
14.5
14
26
25
Temperature (C)
Fig. 10. Spatial distribution of temperature (C) at 10 m depth during January 2002. Temperature estimated by CTD.
0.14
0.12
Specific absorption coefficient by phytoplankton
0.10
a*ph at 440 and 674 nm
0.08
0.06
0.04
0.00
0.065 0.070 0.075 0.080 0.085 0.090 0.095 0.100 0.105
Fig. 11. Linear correlation between the specific absorption coefficient of phytoplankton (a* ph at 440 nm and 674 nm;
m 2(mg Chl-a)1) and zeaxanthin (mg m3) for samples taken at 10m depth during January 2002. The line represents the
tendency at 95% confidence between both variables.
Nez et al. (2004) but lower for those reported for No- to those obtained by Sosik and Mitchell (1995) for the
vember 2002 by Barocio-Len et al. (2006) as a conse- California Current and by Wu et al. (2007) for the north-
quence of changes in the oceanographic conditions after ern South China Sea (1.93.9) during two cruise surveys
La Nia began to wane. In this study the blue/red peak in spring (May) 2001 and late autumn (November) 2002.
ratios ranged from 1.70 to 4.45. These values are similar According to these authors a blue/red ratio number greater
Specific Absorption Coefficient and Phytoplankton Community in the Southern California Current 729
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