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Abstract


A time series was carried out off the coast of Falmouth over an 8 hour period. Using a CTD, ADCP and chemical analysis, the physiological and organic properties of the water column were studied. A thermocline was a clear feature of the water column, and a high Richardson number around such a thermocline further proved the water column was heavily stratified. Nutrients were found in greater abundance around the thermocline, and coupled with that, a higher level of fluorescence, and thus phytoplankton. To act upon the large levels of phytoplankton, a wide diversity of Zooplankton orders were found below the thermocline.


Introduction


During the spring, due to solar heating and weak current the surface layers of the sea will become stratified in offshore Falmouth (Larsonneur et al., 1982). This layer of hotter water will be nutrient rich due to winter upwelling and will normally lead to blooms of diatoms (Menesguen and Hoch, 1997). These blooms in spring will deplete nutrients in the surface layer and die off due to lack of available nutrients (Menesguen and Hoch, 1997). Therefore by mid-summer the surface layer is stratified, warm and nutrient depleted. A thermocline below this surface layer is set up and creates a boundary that restricts nutrients entering the surface waters.


The aim of this study was to assess the effects, indirect or direct if any, of vertical mixing on the temporal variability of the plankton communities and their structure by looking at the behaviour of nutrients throughout the water column. As well as this we wanted to look at the changes mentioned throughout the day and show their variability within the tidal cycle.


Irigoien et al. (2004) demonstrated that the micro- and meso-zooplankton biomass (the latter only detected in the English Channel) saturate with increasing phytoplankton biomass globally. A correlation between strong thermocline and high zooplankton biomass in Western English Channel was documented by Robinson and Hunt (1986). Therefore our hypothesis was that below the thermocline we would find zooplankton maximums as well as the phytoplankton maximum being above but near to the thermocline.


Methods


Rosette Sampler

We used a rosette sampler equipped with 8 Niskin bottles and a CTD, which recorded temperature, salinity and fluorescence (to measure chlorophyll concentration). It was lowered to roughly 5m above the sea floor at each station and the CTD profile of the descent was used to determine where to collect samples during the ascent. Three samples were taken using the Niskin bottles at each location with one near the sea floor, one during the chlorophyll maximum and one near the surface – the collection of which was triggered remotely from the boat.


Plankton net

A plankton net (mesh size 200µm, diameter 0.5m) was deployed at each station twice (three times at the first station) at different depths. The depths were determined based on areas of interest identified on the CTD profile (at the chlorophyll maximum and another below where zooplankton are likely to conjugate).

Laboratory – Callista

Samples taken from the plankton nets and the Niskin bottles were processed and preserved in the on-board laboratory. From the Niskin bottles, separate samples were taken to analyse phytoplankton, dissolved oxygen, phosphate, nitrate, silicon and chlorophyll concentrations. The zooplankton catch was bottled along with a preservative (formalin).


Laboratory – Falmouth

In the laboratory, phytoplankton and zooplankton samples were counted and identified using light and stereo microscopes respectively. These values were then converted to give an average concentration of plankton at each recorded depth in every station.


The water samples were then analysed to give us readings for O2, phosphate, nitrate, silicon and chlorophyll.



Results


Figures 1-12


Temperature

Throughout the tidal cycle from 08:30-16:30 UTC a clear thermocline is visible with a constant temperature range from 12-18oC throughout the observed part of the tidal cycle.


At 08:30 the thermocline initially ranges from 15-20m (Figure 1), dropping to 18-25m by 10:30 (Figure 2). A small pocket of cooler water (13oC) at 18m formed within the thermocline at 09:30 (Figure 4), disappearing by 10:30.


Initially starting at ~10m the thermocline widened at 11:30 (Figure 5) to a depth of ~25m; however this has disappeared by low tide at 12:30 (Figure 6); where the thermocline descended to ~20-30m, before rising to ~15m at 14:30 (Figure 7). 15:30 (Figure 8) saw a lowering of the thermocline to 20-30m, rising slightly by 16:30 (Figure 9) to 18-23m.

As the tide ebbed, a descent of the thermocline was observed, with a subsequent ascent initialising at 14:30 following low tide at 13:30 (Figure 3). The descent of the thermocline coincided with improved mixing of the water column below the thermocline; the temperature remaining at a constant 13oC from 25-60m.


Salinity

Whilst no halocline is present (Figure 10), a clear temporary decrease in salinity can be observed throughout the time series for the duration of the thermocline (Figure 1,2,4,5,6,7,8,9). Two areas of much higher salinity are visible at 09:30 where salinity rises to 36.2 from 35.0, and at 15:30 where a smaller increase from 35.0 to 35.4 is observed. The former spike in salinity coincides with that observed in temperature and density, whilst the latter is only clearly visible in salinity. However it must be noted that the beginnings of a similar small area of great increase are visible in both density and temperature (Figure 3,11).


Density

The pycnocline maintained a similar structure to the thermocline throughout the tidal cycle (Figure 11). Water density ranged from 1025-1026kg/m3 at 0~25m, 1025.5kg/m3 for the duration of the pycnocline and 1026-1027.5kg/m3 below to 60m. Identical to the thermocline, a small pocket of high density water (1027.kg/m3) is visible at 09:30, disappearing by 10:30. However there is a less noticeable descent of the pycnocline to ~25m at low tide (13:30 UTC), before ascending to ~20m where it remains relatively constant.


Slight stratification below the pycnocline is visible. ~25-40m maintained a density of 1026.5-1026.8, increasing to 1027-1027.5 kg/m3 from ~40-60m


Fluorescence

Fluorescence has been used as a proxy for chlorophyll and thereby phytoplankton growth.

Highest fluorescence levels can be observed just below the thermocline at the chlorophyll maximum (Figure 12).


From the surface to the chlorophyll maximum little fluorescence occurred throughout the time series, with the lowest levels (-0.2-0 mg/m3) at the surface.


At 13:30, low tide, fluorescence reduced at depth to 0.08, occurring only at the chlorophyll maximum, for the width of the thermocline (Figure 6). Prior to this a second peak in fluorescence (0.16mg/m3) was visible at ~40m (all multiline plots from 08:30-13:30). However this does not reappear after low tide (all multiline plots from 13:30-16:30), maintaining a constant fluorescence of 0.08-0.10mg/m3 from ~30-60m.


Figures 13-17


Chlorophyll

Chlorophyll follows the same structure as fluorescence, (Figure 13) with low levels in surface waters, increasing from ~0.6mg/L to 1.5-2.5mg/L by 30m, before decreasing to ~1.0 mg/L at 60m.


Between 08:30 to 13:30, a second peak in chlorophyll can be identified at ~45m where chlorophyll rises to a maximum of 3.2mg/l at 15:30 before returning to its steady level of 1.0-1.5mg/L for the remainder of the tidal cycle


O2 (%)

Few changes in dissolved oxygen occur during the time series (Figure 14). At the surface dissolved oxygen ranged from 103-106%, decreasing to 90-97% at 60m. Low tide at 13:30 showed a much steeper gradient, with dissolved oxygen decreasing from 104% to 96% by the thermocline at 25m, before dropping further to 92% at 50m.


Silicate

For the majority of the time series, dissolved silica remained between 0-1 μmol/L from the surface to 30m (Figure 15), before increasing to 1.5-3 μmol/L at 60m. However, at 1130 and 1330, initial surface silica concentrations were much higher (5.8 μmol/L), dropping to 0.7 μmol/L (11:30 UTC) and 1.1 μmol/L (13:30 UTC) by 25m, before increasing back to 1.4-1.5 μmol/L at 50m.


Nitrogen

From the surface to ~25m nitrate concentrations remain relatively constant between 0-1 μmol/L (Figure 16), dropping slightly at 25m to -0.1-0.5 μmol/L, before increasing to range between 0.5-1.5 μmol/L at 50-60m.

At low tide (13:30 UTC) nitrate concentrations were a constant 1.6 μmol/L between 0-25m, before decreasing to 0 μmol/L at 50m.


Phosphate

Across the tidal cycle, phosphate is constantly changing (Figure 17). In general at the surface, concentrations vary between 0-0.1 μmol/L before increasing to concentrations ranging from 0.06-0.45 μmol/L at the thermocline.


At the thermocline a gradual increase in concentration can be observed until low tide, starting at 0.05 μmol/L at 08:30 UTC, reaching 0.25 μmol/L by 10:30, peaking at 11:30 (0.45 μmol/L) before dropping at low tide (13:30) to 0.09 μmol/L, where the decrease continues to 14:30 (0.15 μmol/L) before increasing to a final concentration of 0.38 μ mol/L.


There is no visible pattern at depth. Phosphate concentrations began at -0.1 μmol/L at 65m, rising to 0.15 μmol/L at 65m by 09:30. Subsequently a decrease in concentration to ~0.1 μmol/L before increasing to 0.27 μmol/L at 60m (11:30 UTC) is visible. Low tide saw a drop in concentration to 0.12 μmol/L before rising to 0.38 μmol/L at 16:30.


Figures 18-21


Water Column Structure

Due to lack of data for 08:30-11:30 UTC, Richardson numbers were only able to be calculated from 13:30 to 16:30 UTC.


From the calculated Richardson numbers, it can be seen that the majority of the water column remains well mixed (Figures 18, 19, 20, 21). From low tide at 13:30 to 16:30 UTC, a layer of stratification remains present at 30m. This is most prominent at 13:30 and 16:30, reducing to ~10 from ~30 at 13:30 and increasing to ~60 at 16:30.


A second layer of stratification (Ri of ~40) is present at 14:30 at a depth of 45m. This is visible at all other times, however is far less prominent, with Richardson numbers of 2-5.


A final layer of stratification is visible at ~55m. Whilst this remains throughout the time series, the depth varies from ~58m at 13:30 to a double layer at 53 and 58m at 14:30. The stratification layer remains at 53m at 15:30 before dropping to 55m at 16:30.


Figures 22-27


Phytoplankton

The dominant genera of phytoplankton at the offshore station were Chaetoceros, Guinardia and Leptocylindrus, with Leptocylindrus being present in notably larger quantities than the other genera at all but the first sample at 13:30 UTC (Figure 22). At 15:30 UTC (Figure 23) there was a more consistent spread of plankton between the different genera while all other times show the 3 dominant genera outnumbering the others.

When the data is broken up by depth, very little plankton were present near the surface at the beginning (13:30 UTC) and end (16:30 UTC) of the afternoon time series (Figures 24, 25)) despite these samples having a much higher number of total plankton than the two middle samples (at 14:30 and 15:30 UTC). The two middle samples had a more consistent spread of plankton numbers across all 3 depths. The most dominant genus (Leptocylindrus) generally preferred lower depths of between 25 and 55m and was only found in substantial numbers on the surface at the 15:30 UTC sample.


Figures 28-31


Zooplankton

Figure 28 shows the numbers of different zooplankton orders at station 41 at depths 0-10 metres, 20-30 metres and 30-45 metres, the location of our plankton net samples. What can be seen is that at a depth of 30-45 meters, there is the most diversity on zooplankton orders, however, at a depth of 0-10 metres, they have less species diversity, the same number of orders as is present at 20-30 metres, however a large number of Cladocera were recorded, calculated to be approximately 6367 Cladocera per m-3.


Following station 41, only two plankton nets were sampled at the latter stations, one near the surface and one under a recorded high fluorescence (recorded on CTD data). Figure 29 shows the plankton nets at station 42. Similar to station 1, the deeper depths had greater diversity of zooplankton orders, with 6 at depths 15-25metres, but only 5 at depths of 0-10. Like station 41, there are a great number of Cladocera at a depth of 0-10 metres, with a total calculated number of 12806 Cladocera per m-3.


A much similar trend to the previous stations can be seen at station 43 (Figure 30). Cladocera dominates the water column at a depths of 0-30, with predicted numbers of 2479 per m-3, whereas a large diversity is present at a depths 20-40, that of 7 orders. What can be seen due to the overlap of plankton nets, species which are present at 30-40 metres, and 0-20 metres, as those which are not present in both can be assumed to be found outside of the area of overlap (20-30 metres). Such orders consist of Decapoda larvae, Polychaeta larvae and Gastropod larvae, which may only dwell below 30 metres.


Such a trend continues at the last station sampled, station 44 (Figure 31), However, was can be noted is that Cladocera are no longer the most prevalent order present close to surface. In the plankton net 0-10 metres, more Hydromedusae were counted, with a calculated numbers of 4228 hydromedusae per m-3. The depth 25-40 metres- consistent with the trend, has more order diversity than that of 0-10 metres.


Figures 32-34


ADCP

The change in tides can clearly be seen in the ADCP velocity magnitude (Figure 32) and ship track profile (Figure 33). The turn of the tide occurred at 14:30 UTC, following low tide at 13:30 UTC halfway through the afternoon time series. Note no data is currently available for the morning time series due to it being completed by another group.


Before and after the change in tidal state, areas of higher velocity are present. At the change in tide, velocity drops completely, before reversing direction and increasing back to original levels of 0.3-0.4 m/s.


ADCP can also be used to observe zooplankton and other objects in the water column, such as shoals of fish. Throughout the transect, a persistent layer of zooplankton was present at ~35m, just below the thermocline (Figure 34, although this did decline slightly when the tide turned at 14:30 UTC.


Approximately at 15:30-16:30 UTC, a high area of backscatter ~80dB is visible in the first 10m, predominantly at ~7m. In comparison, the area between the thermocline and the surface contained low backscatter levels ~65-70dB, indicating that there was little present in this area.


Discussion


From the temperature contour it can be seen that the depth of the thermocline increases as the tide goes out. The tide height reaches a minimum of 0.8m at 12:25 UTC. At 08:30 UTC the bottom of the thermocline sits at approximately 21m and this depth decreases as the tide moves out, reaching a depth of approximately 30m at 13:30 UTC. The phytoplankton move with the thermocline so the increase in the depth of the thermocline causes the depth of the chlorophyll maxima to increase.


There is no halocline present in the data, which shows that the water column is well mixed. The spikes in slight increases in salinity seen around the thermocline are due to the CTD measuring salinity from conductivity, causing an error in the measurement until the CTD corrects itself. This is due to lag from the temperature measurements, which causes CTD to constantly correct itself as the CTD descends past a strong thermocline.


After 13:30 UTC the tide begins to flood, from the Richardson numbers it can be deduced that the water column becomes more stratified as the tide comes in. The temperature of the surface water increases due to solar energy. Between 15-25 metres below the surface, a strong thermocline is present. This thermocline drops from approximately 17.5˚C to 12˚C. This becomes more pronounced throughout the day, as the surface waters become heated by solar energy (Paulson & Simpson, 1977).  The thermocline can be seen to be at its deepest at approximately 12:00-13:00 UTC, when the Sun’s rays are at their most powerful.


A small pocket of high salinity, low temperature water can be seen at 09:00-09:30 UTC at a depth of 15-17 metres. This pocket could be caused by a technological error, and poor measurement by the CTD.


Fluorescence maxima (indicative of chlorophyll maximum) can be seen below the thermocline throughout the time series. Phytoplankton typically dwell just above the thermocline area due to an increase in available nutrient. Excluding some cases of silicate, which are higher near the surface, nutrient levels typically increase below the thermocline. This is caused by blooms in the upper water column in earlier months, which deplete the nutrient levels in the upper water column, and thus- to survive- the phytoplankton will migrate downward.


Needing sunlight to photosynthesise, phytoplankton will remain at a depth at which sufficient light and nutrients are available. Phytoplankton thus thrive just above the thermocline, where there is more nutrients than at the surface waters, and enough sunlight to photosynthesise.  These nutrients are supplied by turbulence around the thermocline, and thus providing bottom layer nutrients for production (Sharples et al, 2001). Due to its unique physiological features, there are a vast amount of phytoplankton above the thermocline, so much so that they contribute 20-30% of total primary production in the water column (Revelante & Gilmartin, 1973). Due to this large concentration of phytoplankton, zooplankton focus on  the area for sustenance. Without the need for sunlight and the need to hide from predation, zooplankton are found in great numbers below the thermocline


If the water is deeper, it is seen that there is a greater diversity of zooplankton. This can be seen in all four figures (Figures 27-30), which show a greater number of orders of zooplankton. Zooplankton tend to dwell below levels of chlorophyll maximum- and thus phytoplankton maximum - and typically below the thermocline. With zooplankton being a primary food source of marine organisms, the reduced light levels due to deeper waters makes them harder to be preyed upon, (Pingree et al, 1982). Because of this, the zooplankton dwells below the chlorophyll maximum, which is typically of a depth of 25m (Can be seen on CTD figures beforehand), and are partially protected from predation.

In the upper water column Cladocera are a very prominent genus. It is well documented that where there is a large amount of phytoplankton, there is typically a large amount of Cladocera, with Cladocera being one of the most common zooplankton to be encountered (Frey & David, 1987). It is likely competition which drives the Cladocera to the surface water, surviving against predation by their sheer numbers (Stibor & Herwig, 1992).

The ADCP shows a tidal change at approximately 14:30 UTC, This could account for lower nutrient levels in the water column past 14:30 UTC. The nutrient richer estuarine waters begin moving inland, rather than moving out with the tide to supply the coastal waters with nutrients, and in its stead is the nutrient poor waters from further away from shore. Because of this, the nutrient levels decrease significantly.


References


Frey, & David, G. (1987).The taxonomy and biogeography of the Cladocera. Cladocera. Springer Netherlands, 5 - 17 pp.

Irigoien, X., Huisman, J., & Harris, R. P. (2004). Global biodiversity patterns of marine phytoplankton and zooplankton. Nature429(6994), 863 – 867 pp.

Larsonneur, C., Bouysse, P., & Auffret, J. P. (1982). The superficial sediments of the English Channel and its western approaches. Sedimentology, 29(6), 851 – 864 pp.

Menesguen, A., & Hoch, T. (1997). Modelling the biogeochemical cycles of elements limiting primary production in the English Channel. I. Role of thermohaline stratification. Marine Ecology Progress Series146(1), 173 – 188 pp.

Paulson, C. A., Simpson, J. J., (1977) Irradiance Measurements in the Upper Ocean. J. Phys. Oceanogr., 7, 952 – 956 pp.

Pingree, R. D., et al. (1982) Vertical distribution of plankton in the Skagerrak in relation to doming of the seasonal thermocline. Continental Shelf Research 1(2), 209 - 219 pp.

Revelante, N. and Gilmartin, M. (1973) Some observations on the chlorophyll maximum and primary production in the Eastern North Pacific. Internationale Revue der gesamten Hydrobiologie und Hydrographie. 58(6), 819 – 834 pp.

Robinson, G. A., & Hunt, H. G. (1986). Continuous plankton records: annual fluctuations of the plankton in the western English Channel, 1958–83. Journal of the Marine Biological Association of the United Kingdom66(04), 791 – 802 pp.

Sharples, J., Mark Moore, C., Rippeth, T. P., Holligan, P. M., Hydes, D. J., Fisher, N. R. and Simpson, J. H. (2001) Phytoplankton distribution and survival in the thermoline. Limnol. Oceanogr., 46(3), 486 – 496 pp.

Stibor, & Herwig. (1992) Predator induced life-history shifts in a freshwater cladoceran. Oecologia 92(2), 162 – 165 pp.


Meta Data

Date: 28/06/2014
Time: 13.25

Location: 50°05’610”N 004°52’961”W

Low Tide: 12.25 (0.8m)

High Tide: 05.42 (4.8m), 17.56 (5.0m)

Light Wind

8 octants cloud cover (light rain) reducing to 2 octants

(All times in UTC)


Offshore

Image 1 | CTD rosette pre-deployment.

Image 2 | Cleaning of the zooplankton net.

Disclaimer- the views shown here are solely of group 10 and do not necessarily reflect the views or opinions of the university of Southampton.

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