Marine Microbial Processes

Outline
Part I

A brief history of the development of our current understanding of the
abundance and function of microbes in marine food webs

Part II

The role of microbial processes in carbon (and nitrogen) cycling within
marine food webs

But First, Some Definitions...

Oligotrophic:

Pelagic environment (water column) that has naturally very low plant
nutrient concentrations

--the vast subtropical gyres are oligotrophic

!

Eutrophic:

Pelagic environment (water column) that has naturally high plant nutrient
concentrations

--coastal upwelling zones are eutrophic

As a general rule, the preferred prey size is approximately

1/10 of consumer size

Other than chlorophyll, size determines almost everything about
an organisms role in the community of pelagic organisms

1. It determines who it will eat (all organisms 1/10 its own size)

2. It determines who will eat it (all organisms 10 times bigger than it)

Marine Food Webs are Said to be Strongly Size-Structured!

Simplified Pelagic Food Chain Conceptualization

Optimal Prey Size (1:10) of Pelagic Animals is a Valid General Rule for
Even the Smallest Pelagic Organisms (Protozoans and Bacteria)

Traditional Food
Chain Concept
(early1970s)

Note: organisms are placed into a

given Trophic Level based on
Autotroph versus Heterotrophic
(i.e., chlorophyll versus no
chlorophyll) and on the organisms
size

-----> het. flagellates?

-----> het. bacteria?

auto. bacteria?

Traditional Bacterial
Concentrations
Estimated from
Transmission Light
Microscopy and
Culture-Plate Colony
Counts

Use of Epifluorescent Microscopy and Fluorescent DNA Stains

Became Widespread Between 1975 and 1985

Dramatically increased
estimates of bacterial
concentrations in the ocean

Also allowed easy
distinction between
autotrophic and
heterotrophic flagellate cells
(i.e., chlorophyll containing or
chlorophyll lacking)

10

Bacterial Concentrations
Before (Red Fill) and After
(Blue Fill) the Introduction
of Epifluorescent
Microscopy

11

New view of marine food webs

recognizes the importance of high
bacterial biomass and a large
fraction of nanoflagellates (2-20micron diameter cells) that are
heterotrophic consumers of
bacteria
An unanswered question
What is the carbon and energy source
for all this newly discovered
heterotrophic bacteria?

12

Heterotrophic bacteria are

growing on dissolved
organic matter (sugars,
amino acids etc) released
from phytoplankton by
steady leakage,sudden cell
senescence or sloppy
feeding by zooplankton

13

The term Microbial

Loop is coined by Azam
et al. (1983) to describe
the role that microbes play
in the marine ecosystem
carbon cycle

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Summary: Early 1970s versus Early 1980s

15

Discovery of an Important New

Bacteria-Sized Autotroph
In 1988 Sally Chisholm and Others Published a Paper Describing
the Presence of a New Type of Very Small Autotroph that is
Present in High Abundance - Especially in Oligotrophic Regions

The Discovery was Made using a New Technique called Analytical
Flow Cytometry

This Important New Autotroph Came to be Known as

Prochlorococcus

16

Simple Diagram of Flow

Cytometric Method

17

Prochlorococcus Abundance
1. Prochlorococcus abundance in the oligotrophic open-ocean
is similar in magnitude to the abundance of heterotrophic
bacteria

2. Roughly 1/3 of all bacteria in the oligotrophic open-ocean
is autotrophic Prochlorococcus

18

New View (1990s) of

Marine Food Webs
that Recognizes the
Importance of
Prochlorococcus

19

In oligotrophic (low nutrient) open-ocean environments, the growth

advantage goes to the smallest phytoplankton cells which are now
recognized to be represented mainly by Prochlorococcus.
Prochlorococcus is the main contributor
to primary production in open-ocean
environments.

Altogether, Prochlorococcus singlehandedly contributes more than a
quarter of total ocean primary
production with the remainder being
contributed by hundreds of other
phytoplankton groups.

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Relative Importance of Prochlorococcus and Heterotrophic Bacteria in

Oligotrophic Systems

21

Conclusions
1. Heterotrophic bacteria are highly abundant in all ocean environments

Dissolved organic carbon that leaks from large phytoplankton cells will
be consumed by heterotrophic bacteria and, with the help of protozoans,
this organic carbon will eventually be respired back to carbon dioxide.

2. Prochlorococcus is an autotrophic bacterium (i.e., bacteria that contains
chlorophyll and the smallest type of phytoplankton)

It is the main primary producer in oligotrophic (low nutrient)
environments

It is responsible for more than a quarter of the global ocean primary
production

3. The vast majority of living biomass in the open-ocean is in the form of
heterotrophic bacteria and Prochlorococcus and this was only fairly recently
discovered.

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Marine Microbial Processes II

Carbon & Nitrogen Cycling Within Marine Food Webs

23

Recall From Last Lecture...

New View (1990s) of Marine Food Webs that Recognizes the Importance of
heterotrophic bacteria and autotrophic bacteria (Prochlorococcus)

1970s

1990s

24

Recall...
Oligotrophic:

Pelagic environment (water column) that has naturally very low plant
nutrient concentrations

--the vast subtropical gyres are oligotrophic

!

Eutrophic:

Pelagic environment (water column) that has naturally high plant nutrient
concentrations

--coastal upwelling zones are eutrophic

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The Role of Microbes in Material Flow Through

Marine Ecosystems

26

Recall: Global ocean primary production is an important component of the

global carbon cycle. It is, therefore, an essential part to our understanding of our
global climate and ability predict future global climate change

Biological
Carbon
Pump

Magnitude of CO2 flux between Land and Ocean Reservoirs

27

Recall that low nutrient concentration shifts the growth advantage

to small phytoplankton cell (mainly Prochlorococcus)

28

The pathway that carbon takes from CO2(gas) to particulate organic carbon
(through photosynthesis), and on into particulate organic carbon of higher
trophic levels, varies when nutrient concentration varies
CO2(gas)

CO2(gas)

High Nutrient Conditions ----------------------------------------> Low Nutrient Conditions

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Role of Microbes in Carbon Cycling in

the Ocean
The Biological Carbon Pump

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Carbon Cycling and the Biological Carbon Pump

CO2

CO2

CO2

When the dominant phytoplankton cells are large, the dominant grazers are large and their large fecal
material easily sinks to the deep ocean taking organic carbon down with it - this forms an efficient biological
carbon pump. The opposite is true when the dominant phytoplankton is small and grazers are small and fecal
material is so small it cannot easily sink and the particulate carbon is instead respired back to CO2 and overall
the biological pump is inefficient.

31

Question
What is the expected efficiency of the Biological Carbon
Pump for Costal Upwelling Zones?

(a) The carbon pump is

very efficient in coastal upwelling zones

(b) The carbon pump is
not very efficient in coastal upwelling zones

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Conclusions
As nutrient concentration is reduced, the competitive growth advantage
shifts to small phytoplankton cells
Small phytoplankton cells found at low nutrient concentrations enhances
the percentage of organic carbon that is respired back to carbon dioxide
gas and, consequently, carbon is NOT efficiently pumped into the deep
ocean
Large phytoplankton cells found at high nutrient concentrations increases
the percentage of organic carbon that is pumped to the deep ocean and,
consequently, carbon is efficiently pumped into the deep ocean

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Role of Microbes in Nitrogen Cycling in Marine Ecosystems

1. All living things have a roughly fixed ratio of major elements in their cells
(carbon to nitrogen to phosphorous to oxygen etc..) which is often
referred to a fixed chemical stoichiometry...

2. Because of this fixed chemical stoichometry in all living matter, the
pattern of cycling and export to the deep ocean for all of major
elements will look quite similar. For example...

Where you find carbon being recycled you also find nitrogen being
proportionally recycled

And where you find carbon being exported to the deep ocean you also
find nitrogen being proportionally exported to the deep ocean

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Recall: the Pattern of Carbon Cycling and Export...

CO2

CO2

CO2

When the dominant phytoplankton cells are large, the dominant grazers are large and their large fecal
material easily sinks to the deep ocean taking organic carbon down with it - this forms an efficient biological
carbon pump. The opposite is true when the dominant phytoplankton is small and grazers are small and fecal
material is so small it cannot easily sink and the particulate carbon is instead respired back to CO2 and
overall the biological pump is inefficient.

35

Nitrogen Cycling:

the concept of New & Recycled Primary Production

Total Primary Production = Recycled + New Primary Production

Form of
Nitrogen

Uptake

NH4

1. Recycled Primary
production uses Ammonia
(NH4) generated by
animal excretion in the
upper ocean for its
nitrogen source

2. New Primary
Production uses Nitrate
(NO3) from the deep ocean
for its nitrogen source.

NO3

NH4: Ammonia NO3: Nitrate

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Oligotrophic

Eutrophic

Nitrogen
Uptake

Nitrogen
Uptake

Eutrophic (high nutrients) conditions are dominated by large cell and most of the primary
production is New production

Oligotrophic (low nutrients) conditions are dominated by small cells and most of the
primary production is Recycled production

NOTE: New Production is the portion of total primary production that is available to sink
out of the upper water column and into the deep ocean

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Conclusions
As nutrient concentration is reduced, the competitive growth advantage shifts to small
phytoplankton cells
Oligotrophic (Low Nutrient) Conditions

Small phytoplankton cells and small grazers enhance the percentage of organic carbon that is respired
back to carbon dioxide and carbon is NOT efficiently pumped into the deep ocean (i.e., most carbon
is recycled)

Small phytoplankton cells and microbial grazers and increases the level of nitrogen recycling in the
upper ocean

Eutrophic (High Nutrient) Conditions

Large phytoplankton cells and large grazers increase the percentage of organic carbon that is
pumped to the deep ocean and carbon is efficiently pumped into the deep ocean

Large phytoplankton cells and Large grazers grazers and decreases the level of nitrogen recycling in
the upper ocean

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Evolving Concepts of
Microbial Food Webs

39

High Nutrient Low Chlorophyll Regions (HNLC)

40

Iron Cycling in HNLC

Regions

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Station Aloha - Subtropical North Pacific

Station
Aloha

42

Time Series of N:P Ratio for Total Dissolved, Suspended Particulates

in the North Pacific Subtropical Gyre (from Karl 1999) Showing a shift
to Phosphorous Limitation

43

Phosphorous in the
form of organic
phosphorous or
inorganic phosphate
is taken up by
autotrophs for growth

Phosphorous
Cycling:
organic phosphorous

Inorganic Phosphate

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Nitrogen
Fixation

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Role of Microbes in the Cycling of Other Chemical Elements

Typically oceanographers want to study the cycling of the element
that is limiting the growth of phytoplankton in the region of
interest...

So for the Southern Ocean they study iron cycling

And in the North Pacific Subtropical Gyre (north of Hawaii)
they study the cycling of phosphorous

But for a great many places in the world ocean, nitrogen is
limiting and nitrogen cycling is studied

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