Progress in Oceanography 176 (2019) 102129

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Progress in Oceanography
journal homepage: www.elsevier.com/locate/pocean

Three-dimensional coupling between size-fractionated chlorophyll-a, POC T

and physical processes in the Taiwan Strait in summer
⁎
James T. Liua, , Bangqin Huangb, Yi Changc, Xiaoqin Dua,e, Xin Liub, Rick J. Yanga, Ray T. Hsua,
Saulwood Lind, Jia-Jang Hunga, Jay Leea,f, Chih-Chieh Sud, Yuan-Pin Changa
a
Department of Oceanography, National Sun Yat-sen University, Kaohsiung, Taiwan, ROC
b
Fujian Provincial Key Laboratory of Coastal Ecology and Environmental Studies, College of the Environment and Ecology, Xiamen University, Xiamen, China
c
Department of Hydraulic & Ocean Engineering and Institute of Ocean Technology and Marine Affairs, National Cheng-Kung University, Tainan, Taiwan, ROC
d
Institute of Oceanography, National Taiwan University, Taipei, Taiwan, ROC
e
School of Ocean Science and Technology, Zhejiang Ocean University, Zhoushan, Zhejiang, China
f
Taiwan Ocean Research Institute, National Applied Laboratories, Kaohsiung, Taiwan, ROC

A R T I C LE I N FO A B S T R A C T

Keywords: Coastal water masses are inherently diverse, often with multiple water sources of different origins. Knowledge of
Upwelling characteristic salinity and temperature fingerprints can be very useful to differentiate between component water
River plume masses. For example, summertime river plumes carry relatively warm and less saline water into coastal waters,
Size-fractionated chlorophyll along with nutrients, clastic and organic sediments, and terrestrial-sourced biogenic particles. In an upwelling
Size-fractionated POC
regime, relatively colder water carries marine-sourced nutrients and particles in water masses from deeper
Phytoplankton
C/N ratio
depths offshore. These two regimes are subject to biological-physical coupling that affects their size-differ-
entiated particulate chlorophyll-a and organic carbon contents, and we show here how both regimes can be
identified in these terms in the Taiwan Strait in summer. A close examination shows that microphytoplankton is
the major contributor to chlorophyll-a concentrations in both regimes at all depths, and picophytoplankton is a
minor contributor in the subsurface cold-water regime. The contribution from nanophytoplankton is relatively
insignificant. Shipboard hydrographic profiling and water sampling was conducted over 28 h at a fixed location
off the mouth of the Minjiang River in the northern Taiwan Strait, where the river plume and upwelling regimes
overlap. Water samples were filtered onboard using a nested filtration system to separate suspended particles
into > 153, 63–153, 10–63, and 0.7–10 μm size classes. Our findings show that in the surface nepheloid layer,
the river plume accounted for the physical process affecting large (> 10 μm) terrestrial-sourced non-phyto-
plankton POC (organic debris), with an average C/N ratio of 92.2. The Upwelling regime accounted for the
physical process affecting marine-sourced POC < 10 μm containing fluorescence. In the intermediate nepheloid
layer, which coincided with the subsurface chlorophyll maximum layer, most large (> 10 μm) POC that con-
tained fluorescence were either upwelling- or offshore-sourced, with an average C/N ratio of 8.5. In the benthic
nepheloid layer, terrestrial- and marine-sourced POC containing fluorescence of all sizes were associated with
river plume and upwelling regimes. Large (> 63 μm) non-phytoplankton organic particles were also present,
with an average C/N ratio of 10.2. Both spatial and temporal datasets point to strong three-dimensional coupling
between biogenic particles and physical processes. The river plume and upwelling regimes further differentiate
size-fractionated chlorophyll-a and POC.

1. Introduction
In coastal oceans, nutrient-rich water sustains highly productive
ecosystems that provide fertile fishing grounds. These systems can be
differentiated into two major types, according to the nutrient source
and physical processes by which nutrients are delivered. One type is
associated with coastal upwelling that produces coastal pelagic eco-
systems around the globe (Chang et al., 2014; Chavez and Messié, 2009;
Pérez et al., 2010; Rykaczewski and Checkley, 2008; Wilkerson et al.,
2006). The other type is associated with land-derived nutrients deliv-
ered to the coastal ocean by river plumes (Bainbridge et al., 2012;
Guillaud et al., 2008; Lohrenz et al., 2008; Wysocki et al., 2006).

⁎
Corresponding author.
E-mail address: james@mail.nsysu.edu.tw (J.T. Liu).

https://doi.org/10.1016/j.pocean.2019.102129
Received 9 May 2018; Received in revised form 17 June 2019; Accepted 25 June 2019
Available online 25 June 2019
0079-6611/ © 2019 Published by Elsevier Ltd.
J.T. Liu, et al. Progress in Oceanography 176 (2019) 102129

Because of the dependence on physical processes to deliver nutrients

and provide for environmental stability (e.g. stratification that favors
phytoplankton ecology and growth), these systems exhibit strong bio-
logical and physical coupling that generate variability and stability
(Daly and Smith, 1993; Du and Liu, 2017; McManus and Woodson,
2012; Prairie et al., 2012).
Upwelling and river plume regimes can be observed by ship-based
surveys (Du and Liu, 2017; Lee et al., 2016; Lee Chen, et al., 2004; Liu
et al., 2002; Tang et al., 2002; Valla and Piola, 2015; Wilkerson et al.,
2006). They also can be visualized in satellite images of sea surface
temperature (SST) and chlorophyll/fluorescence (Lee et al., 2015;
Shang et al., 2004, 2011; Tang et al., 2002). The contrast in water mass
composition between the upwelling and river plume regimes, and the
ambient coastal water, often creates surface fronts that can be easily
traced and mapped using satellite-derived information (Chang and
Cornillon, 2015; Jing et al., 2015, 2016; Lan et al., 2009, Jing et al.,
2016). Thermal fronts often mark the boundary of upwelling systems
(Jing et al., 2015, 2016), and can indicate the extent of river plumes as
well. Fronts of chlorophyll-a patches delineate boundaries of high pri-
mary production (Lan et al., 2009). Consequently, biophysical coupling
is manifest in regions where these two types of fronts overlap in the
coastal ocean (Gan et al., 2010; Hong et al., 2009, 2011c; Shang et al.,
2011, Hong et al., 2011c). Since physical processes affect the water
column in horizontal and vertical dimensions, we expect that three-
dimensional structures that reflect biophysical coupling exist, although
they have not been well examined. Furthermore, how the three-di-
Fig. 1. (a) A map showing the area of interest between the South and East
mensional structures affect the size-fractionated biogenic particles and
China Seas, and the Pearl River, a major distal source for fluvial input into the
POC is also worth investigating. Taiwan Strait. (b) Two primary sources of information come from shipboard
In this study, we adopted a straightforward sampling rationale. transects (C, B, A, X1, X4, X1, Y4, Y3, and Y2) and fixed point measurements at
Firstly, we used shipboard surveys along shore-normal transects to the MJ station for shipboard profiling and water sampling in summer, 2012.
create a spatial dataset that covers a wide area with multiple depths Note: The Romanization ‘Zhangyun’ is equivalent to ‘Changyun’ in Taiwan.
that encompasses the two physical regimes. Secondly, we used aerial
data as snap shots of physical changes with built-in space and time could be as high as 2.3 Sv (Hu et al., 2011; Jan et al., 2006). As a
aliasing. To further reveal temporal variability, such as diel biological conduit linking the two major Asian marginal seas, the Taiwan Strait is
activities and tidal influences that snap shots could not resolve, ship- analogous to a monsoon-driven artery intermittently pumping nutrient-
board sampling of multiple depths at a fixed point for 28 h was im- rich water from the upwelling and river plumes that carry biogenic
plemented. A fixed position was chosen at a strategic site where the particles and fluvial sediments into the East China Sea (Azzaro et al.,
influence of both upwelling and river plume regimes could be examined 2007; Xu et al., 2009). Eutrophic water produces patches of high
in detail. chlorophyll-a concentration whose distributions are affected by their
source region and the major topographic barrier imposed by the
2. Study area and background Zhangyun Ridge (Figs. 1 and 2d; Hong et al., 2011c; Shang et al., 2011).
Both river plume and upwelling regimes exist in the Taiwan Strait in
The Taiwan Strait links two major Asian marginal seas: the South summer, and in some regions they overlap (Gan et al., 2009; Han et al.,
China Sea and the East China Sea (Fig. 1). On the eastern side of the 2012; Hu et al., 2011; Tang et al., 2002), creating complexity in the
Taiwan Strait, small mountainous rivers in Taiwan produce dis- associated particle dynamics. Therefore the focus of this study is two-
proportionately high sediment yields (Liu et al., 2013). On the western fold. Firstly, to examine the spatial co-variability of particulate chlor-
side there are additional small rivers that drain into the Taiwan Strait ophyll-a (phytoplankton) and temperature and salinity in the seawater
from the continent (Bi et al., 2015). The Pearl River, a major river in the in the Taiwan Strait (Fig. 1). The purpose is to establish the biophysical
world, is a significant source of riverine nutrients to the Taiwan Strait in coupling that has implications on the dynamics of biogenic particles
summer (Gan et al., 2014; Qu and Kroeze, 2012). (Hong et al., 2011a). Secondly, to investigate the coupling between
Prevailing SW summer monsoon winds along the west coast of the size-fractionated POC containing fluorescence and the physical pro-
Taiwan Strait provide for favorable upwelling conditions (Hu et al., cesses in the surface, intermediate, and benthic nepheloid layers (SNL,
2015). Consequently, several areas in the Taiwan Strait are known to INL, BNL, respectively) off the mouth of the Minjiang River (Fig. 1)
experience upwelling in summer (Hu et al., 2011, 2015; Lan et al., where river plume and upwelling regimes coexist (Fig. 1). A previous
2009; Tang et al., 2002). The combined factors of upwelling, river study shows that the SNL was closely associated with the river plume
plume, and SW monsoon winds in summer, make the Taiwan Strait an regime of the Minjiang River, which was mostly composed of particles
important source for nutrients (especially phosphate) for the East China finer than 10 μm with high fluorescence (Du and Liu, 2017). The INL
Sea (Chung et al., 2001). coincided with the subsurface chlorophyll/fluorescence maximum
The ocean current systems in the Taiwan Strait are highly seasonal. layer, which was located in the pycnocline layer created by insufficient
In the summer the currents flowing northward from the South China mixing from above and below in the water column. The particles in the
Sea to the East China Sea dominate flows in the Taiwan Strait (Hong BNL were sourced from seafloor resuspension and lateral transport (Du
et al., 2011b; Hu et al., 2010, 2011; Jan et al., 2002, 2006). Generally and Liu, 2017). However, the previous study did not distinguish the
these include the extension of the South China Sea Warm Current in the characteristics of particles in terms of size-fractionated POC associated
west and central part of the strait, and the Kuroshio Branch Current with fluorescence.
along the eastern part of the strait (Jan et al., 2002; Hu et al., 2010; For data acquisition on a regional scale, we used a multi-
Hong et al., 2011c). The estimated total northward water transport

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J.T. Liu, et al. Progress in Oceanography 176 (2019) 102129

Fig. 2. Satellite-based maps of sea surface and fluorescence variability and gradients in the Taiwan Strait for July 2012. (a) Sea surface temperature. (b) Sea surface
chlorophyll-a concentration. (c) SST front gradient. (d) Sea surface chlorophyll-a front gradient. Circled numbers are locations of higher chlorophyll-a and lower
temperature values and strong density gradients, including the following: 1. Offshore from the mouth of the Hangjiang River; 2. Offshore of Dongshan Island; 3.
Offshore from the mouth of the Minjiang River; 4. Offshore from Sansha Bay; 5. Over the Taiwan Bank and around the Penghu Islands; 6. Around Zhangyun Ridge; 7.
Offshore from the mouth of the Jiulongjiang River; 8. Along the SW coast of Taiwan.

disciplinary approach employing remotely sensed data from satellite,
and ship-based in-situ data from transect-surveys at multiple depths. We
examined the spatial co-variability among temperature, salinity,
chlorophyll/fluorescence and pigment-based size fraction of micro-,
nano-, and picophytoplankton, and turbidity. The objective was to es-
tablish a three-dimensional framework that accounts for biophysical
coupling. Additionally, shipboard hydrographic profiling and water
sampling were carried out at a location where both upwelling and river
plume are known to exist to investigate depth-related temporal varia-
bility. The fixed-location sampling scheme focused on temporal changes
in size-fractioned POC, associated with fluorescence/phytoplankton or
organic debris, zooplankton, and flocs.
3.2. Shipboard measurements-fixed point mooring at the MJ station
Hydrographic profiling and water sampling was conducted on July
6 and 7, at the MJ station off the mouth of the Minjiang River, in a
water depth of 40 m (Fig. 1), using the R/V Ocean Research II (OR-2) of
the National Taiwan Ocean University (Du and Liu, 2017). The tem-
poral hydrographic profiling dataset were synchronized at 1 m depth
intervals to generate vertical water column structures (Du and Liu,
2017). Water samples were taken from the SNL (3 m below the surface),
INL (at the depth of subsurface fluorescence maximum determined
prior to sampling), and BNL (3 m from the seafloor) at 2 h intervals
between 01:00 July 6 and 03:00 July 7 for suspended particle analysis.

3. Material and methods 3.3. Satellite-based sea surface gradients

3.1. Shipboard measurements and hydrographic surveys along transects
R/V Yanping II (YP-2) of the Fujian Institute of Oceanography,
conducted transect surveys between June 17 and July 7, 2012. At 78
stations along seven shore-normal transects (C, B, A, X4, X1, Y3, Y4,
and Y2) hydrographic profiling of salinity, temperature, fluorescence,
and turbidity were conducted, and water samples were taken for
chlorophyll-a and phytoplankton pigment analysis (Fig. 1). At each
station, 3–6 water samples were taken in the euphotic layer whose
depth varied between 30 and 150 m.
For the month of July 2012, daily images of sea surface temperature
(SST) for the Taiwan Strait acquired by NOAA’s AVHRR (Advanced
Very High-Resolution Radiometer) satellites were collected. Daily
images of chlorophyll concentration of the Taiwan Strait were also
archived using Aqua/MODIS (Moderate Resolution Imaging
Spectroradiometer) ocean color data. SST and chlorophyll frontal pixels
in each daily image were defined statistically by applying the edge
detection approaches of Shimada et al. (2005) and Belkin and O’Reilly
(2009), respectively. This method computed front gradient magnitude
(GM) for both SST and chlorophyll frontal pixels in each image using
the equation (Chang et al., 2008, 2010):
GM = (∂T/ ∂x)2 + (∂T/ ∂y)2 (1)where T denotes SST/chlorophyll
concentration, and the x and y axes are directed towards east and north,

3
J.T. Liu, et al.
respectively. The average SST/chlorophyll gradient was then de-
termined by dividing the number of times the pixel was defined as front
(during July 2012) to composite monthly mean GM front maps (Fig. 2c,
d). The GM scale indicates the intensity of frontal systems. The higher
the GM values, the sharper the boundaries between different water
masses.
3.4. Laboratory analysis of total chlorophyll-a concentration
Seawater (100–500 ml) taken onboard R/V YP-2 was filtered onto
Whatman™ high efficiency microfiber filter GF/F filters of pore size
0.7 μm under low vacuum (< 150 mm Hg). GF/F filters are standard for
Chl-a (chlorophyll-a) measurement (Moran et al., 1999). Afterwards,
the filters were extracted in 95% acetone at −20 °C in the dark for 20 h
and the fluorescence was measured using the non-acidification method
on a Turner-Design fluorometer following Welschmeyer (1994). The
fluorescence was later converted to Chl-a concentration based on a
working curve using a standard (Sigma Chemical Co.). The precision for
the determination was ± 0.02 mg/m3.
3.5. Laboratory analyses of phytoplankton pigment
Seawater (4–8 l, depending on biomass) taken onboard R/V YP-2
was filtered through 47 mm GF/F filters (under a vacuum pressure <
75 mm Hg in dim light) onboard the research vessel, and then was
immediately frozen in liquid nitrogen prior to laboratory analysis (Liu
et al., 2012, 2015). Briefly, phytoplankton pigments were extracted in
dimethylformamide and analyzed using Dionex UltiMate 3000 HPLC
fitted with a 3.5 μm Eclipse XDB C8 column (4.6 × 150 mm, 3.5 μm
particle size, 100 Å pore size, Agilent Technologies, Waldbronn, Ger-
many) with a detector. Pigments were identified using chromatography
with authentic standards (DHI, Denmark) and with diode-array spec-
troscopy (wavelength range: 300–800 nm). Then, the phytoplankton
composition was calculated using the CHEMTAX (Chemotaxonomy)
program, which was built by Mackey et al. (1996) based on phyto-
plankton chemotaxonomy methods. The phytoplankton identified was
divided into three size groups including, microphytoplankton (diatoms
and dinoflagellates); nanophytoplankton (Chrysophytes, Cryptophytes,
Prasinophytes, Prymensiophytes and Chlorophytes); and picophyto-
plankton (Prochlorococcus and Synechococcus). Contributions to the
total Chl-a concentration were estimated from these phytoplankton
groups.
3.6. Analyses of suspended particles
At the MJ station onboard R/V OR-2, water samples were taken
using 10-l X-Niskin bottles mounted on a CTD rosette and were filtered
onboard with a nested Catnet filtration system (Ho et al., 2007; Du and
Liu, 2017; Hsu and Liu, 2010; Lee et al., 2016), that includes three
funnel-shaped Nitex nets of 153, 63, and 10 μm. The funnels were
stacked up in a polycarbonate container (70 × 30 × 30 cm). The 10 μm
net was on the outside of the stack, the 63 μm was in the middle, and
the 153 μm was inside. A 100-ml PE bottle was attached at the end of
each funnel. On board ship, seawater from the Niskin bottles was
drained through tubing into the top of the Catnet filtration system. 1 L
of the filtered (residual) water in the Catnet was collected through a
valve at the bottom of the container. After the seawater in the Catnet
was drained, particles retained on each net were carefully washed into
the PE bottle by hand, using distilled deionized water (DDW). Each
water sample in the PE bottle and in the 1 L residual bottle was further
filtered onto pre-weighed GF/F (0.7 μm) filters using an onboard va-
cuum system. The entire filtration process separated suspended parti-
cles into > 153, 63–153, 10–63, and 0.7–10 μm size classes. The pre-
weighed filters were subsequently stored in a refrigerator at −4 °C for
further analyses. In the laboratory the filters were dried in an oven at
55 °C for 24 h and then weighed again to render the suspended
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Progress in Oceanography 176 (2019) 102129
sediment concentration (SSC). Afterwards POC and PN were quantified
using established procedures (Hung et al., 2000, 2012).
3.7. Empirical Orthogonal/Eigen (EOF) analysis of correlated datasets in
space and time
R/V YP-2 generated two types of spatially correlated datasets. From
the hydrographic survey at each depth four variables of salinity, tem-
perature, fluorescence, and turbidity were produced from CTD sensors.
From the water sample at each sampling depth, variable Chl-a con-
centrations and pigment concentrations of pico-, nano-, and micro-
phytoplankton were analyzed.
R/V OR-2 produced a temporally correlated dataset. At each depth
(surface, mid-depth, and near-bottom), 9 variables were measured
every 2 h. They included temperature, salinity, fluorescence, SSC, size-
fractionated POC > 153, 63–153, 10–63, 0.7–10 μm, and C/N ratio.
Since all the variables were derived from the same water sample/depth,
they were either spatially or temporally correlated and the datasets
were multivariate in nature.
Therefore, EOF was used as the primary analytical tool to decom-
pose spatially demeaned or temporally demeaned correlation (stan-
dardized covariance) into independent/orthogonal modes through
Eigen analysis (Du et al., 2015; Liu et al., 2000). Each eigenmode was
ranked according to the quotient between the eigenvalue of that mode
and the sum of all eigenvalues. The highest mode (Mode 1) explains
most of the co-variability (correlation) in the dataset. Mode 2 explains
less, and so on. In each mode, the variables were grouped according to
the sign (positive or negative) of their eigenvectors. Variables in the
same group co-varied, or varied directly, meaning their data values all
increased or decreased together. Conversely, the variables in an op-
posing group varied inversely. In each mode, the maximum absolute
value of the eigenvector of a variable usually indicates that variable is
the dominant factor affecting the co-variability of that mode. The ei-
genweightings of a particular eigenmode show the spatial or temporal
patterns of that mode. Although EOF is a quantitative tool, the inter-
pretation of EOF results is subjective, depending on the user’s experi-
ence and knowledge of the analyzed variables. In this study, EOF results
were used to link physical processes (represented by salinity and tem-
perature), with biological responses (represented by fluorescence/
chlorophyll and pigment of phytoplankton), and land-sea interaction
(size-fractionated POC and C/N ratio) (Hsu et al., 2014; Lee et al., 2016;
Liu et al., 2002, 2009a, 2009b; Shi et al., 2006).
4. Results
4.1. High spatial co-variability of water properties at multiple depths
The monthly average SST and chlorophyll concentrations for July
2012 show that the spatial variability of these two variables were
highly coupled, being controlled by physical processes and biological
responses (Gan et al., 2010; Hong et al., 2011a) (Fig. 2a, b). The circled
numbers mark six locations of lower temperature (Fig. 2a) con-
ventionally regarded as upwelling regions (Tang et al., 2002). On the
other hand, the seven areas of higher chlorophyll concentration were
mostly located along the coasts of China and Taiwan near river mouths
(#1, 3, 4, 7, 8); in the area south of Penghu Is. (#5); and over the
Zhangyun Ridge (#6, Figs. 1, 2b). The five areas (#1, 2, 3, 4, 5, 6) that
appear in both maps were likely due to upwelling (lower temperature,
higher chlorophyll). There were two regions at Area #1 and 3 where
both upwelling and river plume regimes could overlap. Furthermore,
the upwelling Area #2 had no corresponding higher chlorophyll
(Fig. 2a, b). This might be due to the depletion of nutrients in the up-
welled water (Hu et al., 2015).
Spatial temperature gradients (GM values) were caused by thermal
fronts between water masses of cooler upwelled water and warmer
ambient coastal water (Fig. 2c). These systems lead to higher primary
J.T. Liu, et al.
Fig. 3. Contoured plots of the shipboard measurements of four hydrographic variables at 3, 10, 20, and 30 m depths. (a) Salinity. (b) Temperature. (c) Fluorescence.
(d) Turbidity. Note: the blank space at 30 m is due to the shallow Taiwan Bank (Fig. 1b). The circled numbers are the same as those in Fig. 2.
production as shown by the corresponding areas of higher chlorophyll
(Fig. 2b) that also have sharp gradients (higher GM values) and fronts
(Fig. 2d). Furthermore, the areas of higher chlorophyll that did not
correspond to upwelling were due to the river plume of the Jiulongjiang
River (#7, Yan et al., 2012) and the river plumes on the west coast of
Taiwan (#8). In general, frontal boundaries were marked by sharp
spatial gradients where SST and chlorophyll overlap, and enhance the
coupled nature of the biophysical processes of these systems (Fig. 2c, d).
However, at Area #2 the existence of thermal gradients with the ab-
sence of corresponding chlorophyll gradients suggests depleted nu-
trients in the upwelled water.
One very important variable that indicates the dispersal of river
effluent in the Taiwan Strait is salinity. Based on the shipboard survey
between June-July 2012 (Fig. 1) the spatial salinity structure in the
surface water shows three regions of lower values from Pearl River,
Hangjiang River, Jiulongjiang River, and Minjiang River discharges
(#1, 7, 3, Fig. 3a). Furthermore, the river plume signal can be detected
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Progress in Oceanography 176 (2019) 102129
even at 20 m depth. Outside upwelling areas, the river effluent was also
warmer than ambient Taiwan Strait water throughout the water column
(Fig. 3b). In contrast, the upwelling regime shows lower-temperature
values in the same shipboard data (Fig. 3b). The different sources of
coastal (Area #1, 3, 7) and offshore (Area #5, 6) upwelling regimes are
fairly distinguishable at 20 m depth, separated by a ‘ridge-like’ pattern
of river plume water (Fig. 3b).
Areas of higher chlorophyll concentration as indicated by higher
fluorescence in the shipboard data, largely coincide with the river
plume and upwelling (Fig. 3c). However, one anomalous area offshore
the mouth of the Minjiang River was observed with lower surface
salinity (3 m) and lower temperatures at 3–20 m depths in the ship-
board data (Fig. 3a, b). Since this area also has higher chlorophyll
concentration in the satellite map (area #3 in Fig. 2) and higher
fluorescence at 3–20 m depths in the shipboard data (Fig. 3c), this could
be a region where the upwelling and river plume regimes overlap. To
further clarify this region, shipboard observations were conducted at
J.T. Liu, et al. Progress in Oceanography 176 (2019) 102129

Fig. 4. Contour plot of eigenweightings of Mode 1, 2, 3 at 3 m (a), 10 m (b), and 20 m (c) depths. The percentage values reflect the correlation explained by each
mode. The circled numbers are the same as those in Fig. 2. In each frame, the insert shows the groupings of the 4 variables as indicated by the sign of the eigenvectors.
The contours of eigenweightings show the spatial pattern of the corresponding grouping. The black line is the ‘zero’ contour that separates positive and negative
contoured areas.

the MJ station, off the mouth of the Minjiang River (Fig. 1). At this site,
the shipboard data also show areas of high turbidity (suspended particle
concentration) that do not seem to overlap with salinity, temperature,
and fluorescence patterns (Fig. 3d).
In order to decipher the spatially co-varying nature of the four
variables seen in the shipboard survey, we used EOF analysis at 3, 10,
and 20 m depths (30 m was ignored due to the presence of the shallow
Taiwan Bank).
4.1.1. EOF analysis results at 3 m depth
Near the surface (3 m), Mode 1 explains 39.6% of the correlation,
with temperature and fluorescence in the positive group, and salinity
and turbidity in the negative group according to the eigenvectors
(Fig. 4a, inserts). The two opposing prominent terrestrial and marine
indicators - salinity and temperature, dominate this mode as seen by the
large absolute values of their eigenvectors. The spatial pattern of this
grouping is shown in the contour plot of the eigenweightings, in which
the positive areas follow the coast corresponding to the locations of the
warmer and less salty river plume water (Fig. 3a, b). Therefore, this
mode indicates that the river plume regime (higher temperature, lower
salinity) dominates the hydrographic properties at 3 m depth.
Fluorescence dominates Mode 2 (Fig. 4a, insert), which accounts for
30.5% of the correlation. It co-varies inversely to temperature and
salinity (the lower the temperature and salinity, the higher the fluor-
escence). The negative regions in the contoured eigenweightings of this
mode represent the spatial patterns of weighted fluorescence (Fig. 4a).
Since the shipboard in situ and in vivo fluorescence measurements are
indicative of Chl-a concentration, this mode indicates that high Chl-a

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J.T. Liu, et al.
Table 1
Summary of EOF analysis results of hydrographic vairiables.
Depth Mode 1
3 m (94.0%) 39.6%, temperature (warm river plume)
10 m (94.85) 42.0%, temperature (cold upwelling)
20 m (96.7%) 53.3%, temperature (cold coastal and offshore waters)
Note: The percentage in the parenthesis below the depth is the cumulative percentage of the first 3 modes. The description below each mode percentage is the
dominant variable affecting the co-variability of that mode.
concentration areas are associated with the river plume (lower salinity)
and coastal and offshore upwelling (lower temperature) at 3 m depth.
This region encompasses Areas #1, 2, 3, 5, 6, and 7 shown in Fig. 2.
The third mode explains 23.9% of the data, and is dominated by
turbidity and fluorescence (Fig. 4a, insert) in the negative grouping.
This implies particles with Chl-a, i.e. phytoplankton. The negative areas
in the eigenweighting contours generally overlap with those in the
second mode, i.e. river plume and upwelling regimes, in which the EOF
could not clearly distinguish.
4.1.2. EOF analysis results at 10, and 20 m depths
The eigenstructure (groupings and eigenweighting patterns) for 10,
and 20 m depths are similar to those at 3 m. Therefore, following the
same interpretation principles, the first three modes and their inter-
pretations at all 3 depths (Fig. 4) are summarized in Table 1. These
results show that temperature dominates Mode 1 at all 3 depths
(Fig. 4a, b, c), Mode 2 and 3 are alternately dominated by fluorescence
or turbidity (Fig. 4, Table 1). These structures show the fundamental
control by physical processes, from either river plume or upwelling, on
biological properties (fluorescence), and represent biophysical cou-
pling. The deeper the water, the higher the percentage explained by
Mode 1 (dominated by temperature), and the higher the cumulative
percentage explained by the first three modes (Table 1). This means at
deeper depths, physical processes related to cold-water masses become
more dominant over biological feedback, and increase the co-variability
in the dataset.
4.2. High spatial co-variability in Chl-a and size-fractionated phytoplankton
at multiple depths
To further investigate the spatial co-variability in size-fractionated
biogenic particles and their contribution to Chl-a concentrations cor-
responding to the two major physical regimes in the western Taiwan
Strait, water samples at shipboard transect stations were filtered sepa-
rately both for total Chl-a concentration (Fig. 5a) and for phytoplankton
pigment analyses. The % contributions of tree pigment-based size
groups (micro-, nano-, and picophytoplankton) to the total Chl-a con-
centration were rendered (Fig. 5b-d). Due to various reasons (sampling,
operational, etc.), the filtered dataset has gaps and does not have the
same spatial coverage as the shipboard hydrographic dataset (Figs. 3
and 5). The circled number markings in Fig. 2 are added to Fig. 5 to
facilitate identification of the regime boundaries. The results show that
at all three depths microphytoplankton contributed most to the Chl-a
concentration, followed by pico- and nanophytoplankton, respectively
(Fig. 5b-d). It should be noted that closer to Area #3 in Fig. 1, there was
anomalously higher Chl-a concentration at 3, and 10 m depths (Fig. 5a),
and fluorescence at 3–20 m depths (Fig. 3c). This enhanced primary
production is indicative of overlap between the two regimes (Fig. 3a, b).
EOF analysis was used to decompose spatially correlated tempera-
ture, salinity, turbidity, total Chl-a concentration, and contributions of
the three size-groups at 3, 10, and 20 m depths. The results are plotted
in Fig. 6 and the interpretations are tabulated in Table 2 for clarity.
Since the aim of this study is to understand biophysical coupling, in the
ensuing presentation, the attention is given to biogenic particles that
carry chlorophyll.
7
Progress in Oceanography 176 (2019) 102129
Mode 2 Mode 3
30.5%, fluorescence 23.9%, turbidity
28.9%, fluorescence 23.9%, turbidity
25.6%, turbidity 17.8%, fluorescence
4.2.1. EOF analysis results at 3 m depth
At 3 m depth, Mode 1 (41.8%) shows an association of micro-
phytoplankton, Chl-a concentration, and fluorescence in the negative
group, opposed to temperature (in Fig. 6a, insert). This grouping sug-
gests that microphytoplankton was the major contributor to the Chl-a,
especially in Area #3 (Fig. 5b). This is corroborated by the close re-
semblance of the negative area of the contoured eigenweightings
(Fig. 6a) to the Chl-a concentration patterns (Fig. 5a) and the micro-
phytoplankton distribution at 3 m (Fig. 5b). Also, this group is inversely
related to salinity and temperature, whose eigenweightings overlap
with river plume and upwelling areas (Figs. 3a, b and 6a). Therefore,
this mode reflects a higher microphytoplankton (Figs. 3a,b and 5a,b)
influence in the eutrophic river plume, and coastal and offshore up-
welling regimes (Table 2). In Mode 2 (22.0%) Chl-a, picophyto-
plankton, and turbidity fall in the positive group, and are inversely
related to temperature (Fig. 6a, insert), suggesting an upwelling influ-
ence. On the other hand, because of its weak signal (low absolute ei-
genvector value) the association of fluorescence with other variables is
inconclusive. Mode 2 generally describes the partition of size-fractio-
nated Chl-a being associated with picophytoplankton that was affected
by the coastal upwelling regime. In Mode 3 (12.8%) the salinity and
temperature follow an inverse trend. The turbidity, Chl-a, and pico-
phytoplankton are inversely related to salinity, and nanophytoplankton
is inversely related to temperature. Therefore, Mode 3 describes parti-
tioning between river plume-related terrestrial-sourced particles and
picophytoplankton Chl-a (Fig. 6a, insert).
In summary, the EOF results indicate that river plume and upwel-
ling-associated microphytoplankton dominates Chl-a and fluorescence
distributions at the surface (Mode 1). Upwelling-associated (Mode 2)
and river plume-associated (Mode 3) picophytoplankton is a lesser
contributor to Chl-a. However, Chl-a and fluorescence appeared in
opposed groups in Modes 2 and 3. There are many reasons for the
differentiation between Chl-a and fluorescence by the EOF. Firstly, the
fluorescence signal was a vivo signal, and Chl-a was extracted by or-
ganic solution, after cell breakage. Secondly, Chl-a concentrations ob-
tained for this study were purified Chl-a concentrations, separated with
high performance liquid chromatography/or measured by a Turner
Fluorometer. In addition to Chl-a, there are many other biogenic and
non-biogenic substances that produced fluorescent signals, such as
chromorphic dissolved organic matter. So large differences between
Chl-a and fluorescence signals can occur in coastal waters, especially in
environments with complex physical processes, as in the overlapping
region between river effluent and marine waters.
4.2.2. EOF analysis results at 10 m depth
At 10 m depth, Mode 1 (46.4%) shows an association between mi-
crophytoplankton, fluorescence, Chl-a, turbidity, and salinity in the
negative group, with temperature, nano- and picophytoplankton in the
positive group (Fig. 6b). The negative areas of the contoured eigen-
weightings resemble the cold-water areas at this depth (Fig. 3b).
Therefore, this mode describes size-fractionated Chl-a contributed by
microphytoplankton from the coastal and offshore upwelling regimes.
The Mode 2 groupings (19.3%) are similar to those of the same mode at
3 m depth. But the positive eigenweighting patterns are more similar to
the temperature distribution at 10 m (Figs. 3a and 6b). Thus, this mode
J.T. Liu, et al. Progress in Oceanography 176 (2019) 102129

Fig. 5. Phytoplankton data from filtered seawater samples taken at 3, 10, and 20 m depths. (a) Chl-a concentration; and pigment-based size fraction of (b) mi-
crophytoplankton, (c) nanophytoplankton, and (d) picophytoplankton. The circled numbers are the same as those in Fig. 2. The blank space was caused by lack of
data.

describes partitioning of size-fractionated Chl-a with picophyto-
plankton that was affected by coastal upwelling, with an association of
the other two phytoplankton-groups with the river plume regime. In
Mode 3 (11.6%) the groupings are almost identical to those of the same
mode at 3 m depth, with turbidity having the largest absolute eigen-
vector value. This indicates that terrestrial-sourced turbidity is the
dominant factor affecting this mode. Therefore, the contoured positive
eigenweighting patterns are different (Fig. 6a, b). Generally, this mode
indicates partitioning between river plume-related dispersal of terres-
trial particles and picophytoplankton Chl-a, and upwelling-related
micro- and nanophytoplankton, and fluorescence (Table 2).
In terms of size-fractionated Chl-a at the 10 m depth, the coastal and
offshore upwelling regimes had the largest effect on the Chl-a of the
micro- and picophytoplankton (Modes 1, 2). The river plume regime

8
J.T. Liu, et al. Progress in Oceanography 176 (2019) 102129

Fig. 6. Contour plot of eigenweightings of Mode 1, 2, 3 at 3 m (a), 10 m (b), and 20 m (c) depths. The percentage values reflect the correlation explained by each
mode. The circled numbers are the same as those in Fig. 2. In each frame, the insert shows the groupings of the 8 variables as indicated by the sign of the eigenvectors.
The contours of eigenweightings show the spatial pattern of the corresponding grouping. The black line is the ‘zero’ contour that separates the positive and negative
contoured areas. The blank space was caused by lack of data.

Table 2
Summary of EOF analysis results of size-fractionated phytoplankton and physical regimes.
Depth Mode 1 Mode 2 Mode 3

3 m (76.6%) 41.8%, microphytoplankton/salinity (river plume regime) and 22.0%, picophytoplankton/temperature 12.8%, turbidity and picophytoplankton/
temperature (coastal and offshore upwelling regimes) (coastal upwelling regime) salinity (river plume regime)
10 m (77.3%) 46.4%, microphytoplankton/temperature (coastal and offshore 19.3%, picophytoplankton/temperature 11.6%, turbidity and picophytoplankton/
upwelling regimes) (coastal upwelling regime) salinity (river plume regime)
20 m (79.3%) 48.2%, microphytoplankton/temperature (coastal and offshore 18.1%, microphytoplankton/salinity (river 13.0%, turbidity and nanophytoplankton/no
upwelling regimes) plume regime) dominant physical regimes

Note: The percentage in the parenthesis below the depth is the cumulative percentage of the first 3 modes. The description below each mode percentage is the
phytoplankton group affecting the co-variability of that mode. The dominant physical variable (regime) is indicated after the slash.

9
J.T. Liu, et al.
was a secondary process affecting picophytoplankton Chl-a. (Mode 3)
(Table 2).
4.2.3. EOF analysis results at 20 m depth
At 20 m depth the eigenweighting pattern of Mode 1 (48.2%) was
almost identical to the same mode at 10 m (Fig. 6b, c). Therefore, the
same interpretation applies (Table 2). The temperature and salinity in
Mode 2 (18.1%) fall in opposite groups. The microphytoplankton Chl-a
and turbidity are inversely related to salinity (Fig. 6c), suggesting a
river plume influence. Consequently the contoured patterns of the ne-
gative area of the eigenweightings also resemble those of the salinity
distribution at this depth (Fig. 3a). Therefore, this mode is interpreted
as a river plume influence on microphytoplankton Chl-a (Table 2).
Mode 3 (13.0%) shows that temperature is inversely related to Chl-a,
nano-, picophytoplankton, and turbidity (Fig. 6c). Since turbidity is the
dominant factor, the contoured patterns of the positive eigenweightings
resemble the turbidity distribution (Fig. 3d). Yet, this is the only mode
where nanophytoplankton is found to be associated with fluorescence
(Fig. 6c). Since this mode only explains 13% of the correlations, no
dominant physical regime was inferred (Table 2).
In summary, at 20 m physical processes of the upwelling regime
(coastal and offshore) (Mode 1) and the river plume (Mode 2) were
dominant (Table 2). Microphytoplankton was the primary contributor
to Chl-a (Mode 1, 2). However, when turbidity became the dominant
variable in Mode 3, no clear physical regime could be identified by EOF
analysis, although wave resuspension was speculated. In this mode
only, nanophytoplankton contributed to Chl-a.
4.3. Particles dynamics of size-fractionated POC off the mouth of the
Minjiang River in SNL, INL, and BNL
4.3.1. Difference in water-mass fingerprints in the water column
Temporal changes of all measured variables at 2-h intervals from 6
July 01:00 to 7 July 03:00 show marked salinity and temperature
contrasts between the surface and bottom water (Fig. 7a, b). At the
surface, in the SNL, the Minjiang River plume is clearly seen from low
salinity and high temperature values (Du and Liu, 2017). Although not
fully resolved in the water sample data, measured surface salinity
shows a semi-diurnal tidal influence (Fig. 7a) (Du and Liu, 2017). Near
the bottom, the temperature was about 6 °C lower than at the surface
and salinity was 2–5 PSU higher than that at the surface. Except for
three data points, SSC values at the bottom, in the BNL, were con-
sistently higher than those at the surface and at mid-depth (Fig. 7c).
Due to instrumental errors, the fluorescence values at three time points
were not available (Fig. 7d), however this gap does not hinder the
merits of the dataset as a whole. At a glace, the fluorescence data seems
to have two parts. Before 17:00 on July 6, the fluorescence in the water
column was more or less uniform. However, after this point, the SNL
values increase dramatically, to 2–4 times higher than those in the INL
and BNL (Fig. 7d). This was caused by the presence of nutrient-rich
river plume water in the SNL (Du and Liu, 2017).
4.3.2. C/N ratios and composition of size-fractionated POC in the water
column
For clarity, particle-related data (C/N ratio and size-fractionated
POC) of different depths are presented for the SNL (Fig. 7e, f), INL
(Fig. 7g, h), and BNL (Fig. 7i, j). A quick overview of the data shows
that C/N ratios were the highest at the surface and the lowest near the
bottom (Fig. 7e, g, i). The average C/N values were 92.3 in the SNL, 8.5
in the INL, and 10.2 in the BNL, with standard deviations of 46.0, 4.5,
and 5.1, respectively. The exceptionally high C/N values in the SNL
were likely due to land-sourced woody debris (Yang and Luo, 2011;
Saki et al., 2012). This provides strong evidence for terrestrial particles
sourced from the river effluent along the coast. The total POC con-
centration in the SNL was an order of magnitude higher than those in
the INL and BNL, which were about the same (Fig. 7f, h, j). The < 10
10
Progress in Oceanography 176 (2019) 102129
μm POC size fraction was the largest at all depths, and more dominant
in the INL and BNL. The SNL contained higher proportions of relatively
large POC sizes, particularly 63–153 and > 153 μm (Fig. 7f). These
differences were examined using EOF analysis and are discussed later.
4.3.3. EOF analysis results of particle dynamics in the SNL, INL, and BNL
For each depth, EOF analysis was used to examine the co-variability
among salinity and temperature (water mass indicators), fluorescence
(representing Chl-a and thus phytoplankton), POC > 153, 63–153,
10–63, and < 10 μm (indicating size-fractionated biogenic particles and
organic debris), and C/N ratio (indicator for terrestrial or marine-
sourced POC).
Similar to previous cases, the EOF mode groupings are indicated by
the signs of the eigenvectors (Fig. 8a, b, c). The basic interpretation
scheme is as follows: when the POC is associated with fluorescence (in
the same group), it is interpreted as phytoplankton. Otherwise, the POC
could be zooplankton, organic debris, or floc particles. When POC, SSC,
and C/N were inversely related to salinity, they are interpreted as being
associated with the river plume regime. When they were inversely re-
lated to temperature, they are interpreted as being associated with the
upwelling regime. Alternatively, when the above variables were di-
rectly related to temperature and salinity, they are interpreted as off-
shore-sourced (from water masses in the Taiwan Strait). The higher the
cumulative percentage of correlations explained by the first three ei-
genmodes, the more highly correlated the dataset is.
In the SNL (Fig. 8a), the correlations explained by the first three
modes were 44.2%, 25.2%, and 11%, respectively. The river plume
regime affected POC of all sizes except for POC < 10 μm, SSC, and C/N
(Mode 1); POC 10–63 μm (Mode 2); and SSC, POC > 153, < 10 μm
(Mode 3). The upwelling regime affected fluorescence carried by
POC < 10 μm (Mode 1); POC 10–63 μm (Mode 2); and SSC, POC >
153, < 10 μm (Mode 3). Offshore water masses also affected SSC, C/N
and all POC sizes except for 10–63 μm, that carried fluorescence (Mode
2). In Mode 3, the eigenvalue of fluorescence is too small and therefore
is not associated with any POC size.
In the INL (Fig. 8b), the correlations explained by the first three
modes were 31.6%, 20.4%, and 18.5%, respectively. Both the river
plume and upwelling regimes were resolved in Mode 1. The river plume
regime affected the 10–63 μm POC fraction in the SSC, and the up-
welling regime affected C/N and fluorescence in all POC sizes, except
for 10–63 μm. In Mode 2 the temperature and salinity were in the same
group, pointing to an offshore water mass influence. Also in this mode,
it is inconclusive which regime affected SSC and POC < 10 μm that
were in the same group. Due to low eigenvector values of salinity and
temperature in Mode 3, no physical regime could be identified to ex-
plain the rest of the variables. However, SSC co-varied with fluores-
cence, which was carried by all POC sizes except 63–163 μm. In the
opposite group, C/N was associated with the 63–163 μm POC fraction.
In the BNL (Fig. 8c), the correlations explained by the first three
modes were 46.8%, 24.3%, and 12.3%, respectively. In Mode 1, the
upwelling regime affected C/N and finer POC, 10–63 and < 10 μm, that
carried fluorescence. The river plume only affected POC > 153 μm. In
Mode 2, both temperature and salinity have small eigenvector values.
So their influence is inconclusive. Nevertheless, the upwelling could
affect POC > 153 and 63–153 μm, and C/N values. The river plume
could affect SSC, POC 10–63 and < 10 μm. The eigenvalue of fluores-
cence in this mode is too small to associated with a POC size fraction. In
Mode 3, the river plume affected fluorescence carried by POC > 153,
10–63, and < 10 μm. The upwelling influence is inclusive, due to the
temperature’s small eigenvector value. It could affect SSC, POC 63-153,
and C/N.
J.T. Liu, et al. Progress in Oceanography 176 (2019) 102129

Fig. 7. Hydrography and characteristics of organic particles at three depths observed at the MJ station over 28 h. (a) Salinity, (b) temperature, (c) SSC, and (d)
fluorescence. C/N ratio and cumulative concentrations of POC sizes > 153, 63–153, 10–62, and < 10 μm at 3 m depth (e, f), mid-depth (g, h), and 3 m above seabed
(i, j).

5. Discussion and conclusions
5.1. Limitations to the size determination of phytoplankton and POC
Since the major foci of this study are size-fractionated phyto-
plankton pigments and POC, we need to discuss some constraints in our
methodology. Firstly, based on the chemical classification of pigments,
there are some limitations or uncertainties. Some phytoplankton groups
have specific diagnostic pigments (Wright et al., 1991). Consequently,
results of the HPLC-CHAMTAX method for these groups are more reli-
able. For other groups with low specificity of pigments this method may
have some biases (Antajan et al., 2004). Secondly, the CHEMTAX
method assumes that the ratio of phytoplankton pigments to Chl-a in
one group is relatively fixed. However, many studies have shown that
light, nutrients and other environmental factors can significantly
change the ratios (Jeffrey, 1981), and species within a group can

11
J.T. Liu, et al.
exhibit significant differences in the ratios (Antajan et al., 2004; Zapata
et al., 2004).
Nevertheless, a large number of studies have confirmed that the
results of this method (especially dominant groups) are credible, as
compared with microscopy and flow cytometry. Similar comparisons
and analyses have been made in the vicinity of our study area (Liu et al.,
2016; Xiao et al., 2018). Uncertainties have been discussed and relevant
basis has been provided (Liu et al., 2016; Xiao et al., 2018). Therefore,
for what we want to achieve in this study, we feel confident with the
CHEMTAX method.
The size-fractionated POC, on the other hand, was based on using a
C/N/S analyzer of physically filtered particles using nets and filter
papers of different sizes. Although both types of data have to do with
‘sizes’, they are not strictly comparable. Therefore, we do not make
direct comparisons in our interpretations.
5.2. Three-dimensional coupling on a regional scale due to river plume and
upwelling regimes
On a regional scale, surface water from both the river plume and
coastal upwelling regimes are important physical processes affecting
the dynamics of size-fractionated Chl-a (Tables 1 and 2). Micro-
phytoplankton is associated with both regimes (depicted in Mode 1,
Table 2). This suggests a microphytoplankton-dominated phyto-
plankton structure on a regional scale. This scenario fits the eutrophic
conditions of a river plume (Armbrecht et al., 2015; Van Oostende
et al., 2015; Wilkerson et al., 2006). Picophytoplankton is depicted in
Mode 2 (associated with upwelling) and Mode 3 (river plume) is of
secondary importance. Nanophytoplankton is less important at the
surface, as shown by its smaller contribution to Chl-a concentrations
(Fig. 5c).
At mid-depth, coastal and offshore upwelling regimes became the
dominant physical process (depicted in Mode 1, 2, Table 2) on a re-
gional scale. The river plume was less important (in Mode 3). In this
setting, microphytoplankton was the dominant group (depicted in
Mode 1), picophytoplankton was the secondary group (depicted in
Mode 2, 3). In deeper (20 m) waters, coastal and offshore upwelling
regimes were also the dominant physical process (depicted in Mode 1,
12
Progress in Oceanography 176 (2019) 102129
Fig. 8. EOF analysis results from the MJ
station dataset showing eigenvectors of the
first three eigenmodes for the co-variability
of temperature, salinity, fluorescence, SSC,
POC > 153 μm, POC 63–153 μm, POC
10–63 μm, POC < 10 μm, and C/N at (a) the
surface (3 m), (b) mid-depth, and (c) bottom
(3 m above seabed). Note: eigenvectors in
green colors indicate POC sizes carrying
fluorescence in an eigenmodes. Otherwise,
orange is used. (For interpretation of the
references to color in this figure legend, the
reader is referred to the web version of this
article.)
3). The river plume regime was the second most important physical
process affecting this depth. At this depth, microphytoplankton was the
dominant group (depicted in Mode 1, 2) and picophytoplankton was
the secondary group (in Mode 3). Overall, microphytoplankton was the
dominant group at all depths, followed by picophytoplankton. Nano-
phytoplankton, having the lowest percentage contribution to Chl-a
(Fig. 5c), was only associated with Chl-a at 20 m depth in Mode 3
(Table 2). Thus, they were not an important contributor to the Chl-a
concentration in the study area.
The basic water-mass variables of salinity, temperature, fluores-
cence, and turbidity in the water column were highly correlated. The
total correlation of the first eigenmodes increased with depth from
94.0% to 96.7% (Table 1). Temperature and salinity were the dominant
influencing factors at all three depths, which were controlled by the
interplay between warmer river plume water and colder upwelled
water. This basic eigenstructure remained similar after the addition of
total Chl-a concentration and size-fractionated contribution of Chl-a
(%) in the EOF analysis. The correlation explained by the first three
eigenmodes also increased with depth from 76.6% to 79.3% (Table 2).
In this structure, the river plume and coastal upwelling regimes domi-
nated the physical processes at the surface. The two upwelling regimes
dominated the subsurface water column. However, the additional bio-
geochemical variables in the second typed datasets of 8 variables in-
troduced more variability, which caused the datasets to be less corre-
lated than the hydrographic datasets of 4 variables.
5.3. Temporal co-variability in nepheloid layers off the mouth of the
Minjiang River showing land-sea interaction through size-fractionated POC
The comparable study by Du and Liu (2017) provided background
on the physical processes and particle dynamics of the SNL, INL, and
BNL from which the water samples in this study were taken. At this
location, the tidally modulated dispersal of the river plume was the
dominant physical process affecting the water column stratification at
the surface (Du and Liu, 2017). The river effluent was also the major
source for terrestrial-sourced suspended sediment (SSC) and large POC
sizes (> 10 μm) (Mode 1, Fig. 8a). In fact, non-phytoplankton POC
(organic debris, etc.) dominated the POC structure in the SNL. The
J.T. Liu, et al.
terrestrial origin of this POC was also supported by the high C/N ratios
(Fig. 7e) from woody debris. However, in the SNL upwelling supplied
fine-grained POC with fluorescence (< 10 μm). Both the river effluent
and upwelled water masses carry living and non-living POC of terres-
trial and marine origins in the SNL. Additionally, offshore water masses
also carry terrestrial-sourced SSC and living POC in the SNL, to a lesser
degree (Mode 2, 3). Therefore, there were diverse water masses that
delivered terrestrial and marine-sourced sediment and living and non-
living POC to the SNL.
Because of the co-location of the subsurface chlorophyll maximum
layer and INL (Du and Liu, 2017), the C/N ratio values in the INL were
the closest to that of the Redfield Ratio (Redfield, 1934, 1958) among
the 3 nepheloid-layers. The first 3 eigenmodes of INL only account for
70.5% of the correlations, indicating that they were the least correlated
among the 3 nepheloid layers. Although Mode 1 indicates upwelling
affected the living POC of terrestrial origin and river plume affected SSC
and non-living 10–63 μm POC, it only accounts for 31.6% of the cor-
relation. There were no strong physical processes affecting variables in
the INL, indicated by EOF analysis. Physically, this was because the INL
was formed in the pycnocline layer at mid-depth due to the lack of
vertical mixing of the entire column from the wind above and the
current below (Du and Liu, 2017). Du and Liu (2017) also found that
that in the INL most particles had volume concentration in < 153,
63–153 μm sizes. Our findings corroborate this finding in that most
fluorescence was associated with larger POC > 153, 63–153 μm (Mode
1, 2), and to a lesser degree 10–63 μm (Mode 2) (Fig. 8b). Since the INL
was isolated from the river plume water at the surface and from water
masses near the seabed, the SSC in INL was of non-terrestrial origin (in
opposite groups, Fig. 8b).
Du and Liu (2017) found that particles in the BNL were either re-
suspended off the seabed, or transported laterally by currents, which
could be of both marine and terrestrial origins. Our EOF analysis
findings corroborate the above. We found that in the BNL the upwelling
regime was the most important physical process (in Mode 1) that af-
fected living POC of terrestrial origin (Fig. 8c). The river plume was of
lesser importance (in Mode 3) to bringing living POC of marine origin
to the BNL. However, the SSC of marine origin in Mode 2 was neither
related to upwelling nor river plume, which could be due to re-
suspension.
Overall, the eigenstructure of the temporal dataset showed some
discrepancy. There were direct physical processes affecting the SNL and
BNL, leading to highly correlated datasets, having 80.4% and 83.4% of
correlations explained by the first 3 eigenmodes, respectively. Contrary
to the above, the lack of direct influence of physical processes caused
the INL dataset to be less correlated (70.5%). In other words, strong
coupling between physical processes and biogeochemical responses will
create high co-variability among the studied variables.
5.4. A cross-disciplinary study examining spatial and temporal co-
variability in the coupling between physics and biogeochemistry
Findings from the spatial variability of satellite-derived images,
shipboard transect surveys, and fixed-point coverage of the temporal
variability in particle dynamics complement one another. They point
out the general theme in summer, that along the west coast and the
central part of the Taiwan Strait warm river plume water and cold
subsurface water of upwelling regimes affect salinity and temperature
structures in a three-dimensional fashion. These structures in turn in-
fluence the properties of size-fractionated POC, Chl-a, and suspended
sediment. Biogenic particles were analyzed as the percent contribution
to the Chl-a by micro-, nano-, and picophytoplankton, based on size-
fractionated pigment in the regional-scaled dataset. In our temporal
dataset, complex interactions between river plume-induced stratifica-
tion and wind- and current-induced mixing in the water column directly
controlled the co-variability of measured variables in the three NLs. The
variables were also affected by the water masses that carried them (Du
13
Progress in Oceanography 176 (2019) 102129
and Liu, 2017).
This research is a cross-disciplinary study with a broad range of
interests. The complex biogeochemical coupling with physical pro-
cesses (upwelling and river plume) in coastal waters is a very important
and basic research subject. Importantly, the signals from river effluent
and upwelled waters are different. Rivers carry terrigenous inputs on a
regional scale, in which the effects of human activities (nutrient con-
centrations and their ratios, destruction of forests, etc.) are evident.
Upwelling carries marine-sourced signals, mainly influenced by large-
scale wind and currents. It is very important to distinguish the effects
from both in the coastal environment. They are reflected in the eco-
system dynamics in our study area.
In conclusion, our dataset is comprehensive, with clear process-re-
sponse implications. We used EOF analysis to decipher the complex
associations of co-varying physical, biological, and biochemical vari-
ables. Our study shows that size-fractionated Chl-a concentrations and
size-fractionated POC are highly correlated with the three-dimensional
physical structures of the water column, controlled by the river plume
and coastal and offshore upwelling regimes. Different physical regimes
further differentiate these particles and water masses of different ori-
gins that carried them.
Acknowledgements
This study was supported by the ROC Ministry of Science and
Technology (formerly The National Science Council) under grant
numbers: NSC 100-2611-M-110-013, NSC 101-2611-M-110-013; and
the Natural Science Foundation of China under the grant number NSFC
41330961. Xiamen University hosted JTL’s visit where part of the re-
search was conducted. We are grateful to an anonymous reviewer
whose detailed and helpful comments helped to improve the manu-
script. MRL proofread an earlier version and GB, who proofread a later
version of the manuscript.
Appendix A. Supplementary material
Supplementary data to this article can be found online at https://
doi.org/10.1016/j.pocean.2019.102129.
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