25.2B: Charales - Biology

25.2B: Charales - Biology

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Algae in the order Charales live in fresh water and are often considered the closest-living relatives of embryophytes.

Learning Objectives

  • Identify the principle features of charophyte algae

Key Points

  • The structure of charophyte algae consists of a thallus, which is the main stem, and branches that arise from nodes which bear both male and female reproductive structures.
  • Although charophyte algae do not exhibit alteration of generations, they share a number of adaptations to life on land with embryophytes, including the encasement of eggs in protective enclosures.
  • As new DNA sequence analysis techniques develop, revisions may need to be made in our understanding of plant evolution, such as indications that green algae in the order of Zygnematales may be more-closely related to embryophytes than is Charales.

Key Terms

  • Charales: green algae in the division Charophyta which are green plants believed to be the closest relatives of the green land plants
  • sporopollenin: a combination of biopolymers observed in the tough outer layer of the spore and pollen wall


Green algae in the order Charales, and the coleochaetes, microscopic green algae that enclose their spores in sporopollenin, are considered the closest-living relatives of embryophytes. The Charales can be traced as far back as 420 million years. They live in a range of fresh water habitats and vary in size from as small as a few millimeters to as large as a meter in length. A representative species of Charales is Chara, which is often called muskgrass or skunkweed because of its unpleasant smell.

In Charales, large cells form the thallus: the main stem of the alga. Branches arising from the nodes are made of smaller cells. Male and female reproductive structures are found on the nodes; the sperm have flagella. Unlike land plants, Charales do not undergo alternation of generations in their lifecycle. Like embryophytes, Charales exhibit a number of traits that are significant in their adaptation to land life. They produce the compounds lignin and sporopollenin. They form plasmodesmata, which are microscopic channels that connect the cytoplasm of adjacent cells. The egg and, later, the zygote, form in a protected chamber on the parent plant.

New information from recent, extensive DNA sequence analysis of green algae indicates that the Zygnematales are more closely-related to the embryophytes than the Charales. The Zygnematales include the familiar genus Spirogyra. As techniques in DNA analysis improve and new information on comparative genomics arises, the phylogenetic connections between species will probably continue to change. Clearly, plant biologists have yet to solve the mystery of the origin of land plants.

Comparative analysis of BAC and whole genome shotgun sequences from an Anopheles gambiae region related to Plasmodium encapsulation

The only natural mechanism of malaria transmission in sub-Saharan Africa is the mosquito, generally Anopheles gambiae. Blocking malaria parasite transmission by stopping the development of Plasmodium in the insect vector would provide a useful alternative to the current methods of malaria control. Toward this end, it is important to understand the molecular basis of the malaria parasite refractory phenotype in An. gambiae mosquito strains. We have selected and sequenced six bacterial artificial chromosome (BAC) clones from the Pen-1 region that is the major quantitative trait locus involved in Plasmodium encapsulation. The sequence and the annotation of five overlapping BAC clones plus one adjacent, but not contiguous clone, totaling 585 kb of genomic sequence from the centromeric end of the Pen-1 region of the PEST strain were compared to that of the genome sequence of the same strain produced by the whole genome shotgun technique. This project identified 23 putative mosquito genes plus putative copies of the retrotransposable elements BEL12 and TRANSIBN1_AG in the six BAC clones. Nineteen of the predicted genes are most similar to their Drosophila melanogaster homologs while one is more closely related to vertebrate genes. Comparison of these new BAC sequences plus previously published BAC sequences to the cognate region of the assembled genome sequence identified three retrotransposons present in one sequence version but not the other. One of these elements, Indy, has not been previously described. These observations provide evidence for the recent active transposition of these elements and demonstrate the plasticity of the Anopheles genome. The BAC sequences strongly support the public whole genome shotgun assembly and automatic annotation while also demonstrating the benefit of complementary genome sequences and of human curation. Importantly, the data demonstrate the differences in the genome sequence of an individual mosquito compared to that of a hypothetical, average genome sequence generated by whole genome shotgun assembly.

Cancer cells can alter physiological mechanisms within bone resulting in high bone turnover, and consequently in skeletal-related events (SREs), causing severe morbidity in affected patients. The goals of bone targeted therapy, as bisphosphonates and denosumab, are the reduction of incidence and the delay in occurrence of the SREs, to improve quality of life and pain control.

The toxicity profile is similar between bisphosphonates and denosumab, even if pyrexia, bone pain, arthralgia, renal failure and hypercalcemia are more common with bisphosphonates, while hypocalcemia and toothache are more frequently reported with denosumab. Osteonecrosis of the jaw (ONJ) occurred infrequently without statistically significant difference.

The present review aims to provide an assessment on bone targeted therapies for preventing the occurrence of SREs in bone metastatic breast cancer patients, critically analyzing the evidence available so far on their effectiveness, in light of the different mechanisms of action. Thus, we try to provide tools for the most fitting treatment of bone metastatic breast cancer patients.

We also provide an overview on the usefulness of bone turnover markers in clinical practice and new molecules currently under study for the treatment of bone metastatic disease.

BIOL 181: Life in the Oceans

1. Introduction: Science and Marine Biology 2. Fundamentals of Ecology 3. Marine Provinces 4. Seawater 5. Tides 6. Biological Concepts 7. Marine Microorganisms 8. Multicellular Primary Producers 9. Sponges, Cnidarians, and Comb Jellies 10. Worms, Bryozoans, and Mollusks 11. Arthropods, Echinoderms, and Invertebrate Chordates 12. Marine Fish 13. Marine Reptiles and Birds 14. Marine Mammals 15. Intertidal Ecology 16. Estuaries 17. Coral Reef Communities 18. Continental Shelves and Neritic Zone 19. The Open Ocean 20. Life in the Ocean’s Depths 21. Marine Birds and Mammals in Polar Seas 22. Artificial Reefs 23. Marine Protected Areas 24. Impact of Tourism on the Marine Environment 25. The Global Trade of Marine Ornamental Species

BIOL 181: Life in the Oceans – Lecture Notes

1. Introduction: Science and Marine Biology (The majority of the text below originally appeared as chapter 1 of Introduction to Marine Biology)

1.1. Science and Marine Biology

Oceans cover 71 percent of the earth, and affect climate and weather patterns that in turn impact

the terrestrial environments. They are very important for transportation and as a source of food,

yet are largely unexplored it is commonly said that we know more about the surface of the moon

than we do about the deepest parts of the oceans!

Oceanography is the study of the oceans and their phenomena, and involves sciences such as

biology, chemistry, physics, geology, and meteorology. Marine biology is the study of the

organisms that inhabit the seas and their interactions with each other and their environment.

1.2. Brief History of Marine Biology

Marine biology is a younger science than terrestrial biology, as early scientists were limited in

their study of aquatic organisms by lack of technology to observe and sample them. The Greek

philosopher Aristotle was one of the firsts to design a classification scheme for living organisms,

which he called ―the ladder of life‖ and in which he described 500 species, several of which were

marine. He also studied fish gills and cuttlefish. The Roman naturalist Pliny the Elder published

a 37-volume work called Natural History, which contained several marine species.

Little work on natural history was conducted during the middle ages, and it wasn’t until the late

eighteenth century and early nineteenth century that interest in the marine environment was

renewed, fueled by explorations now made possible by better ships and improved navigation

techniques. In 1831, Darwin set sail for a five-year circumnavigation on the HMS Beagle, and

his observations of organisms during this voyage later led to his elaboration of the theory of

evolution by natural selection. Darwin also developed theories on the formation of atolls, which

turned out to be correct. In the early nineteenth century, the English naturalist Edward Forbes

suggested that no life could survive in the cold, dark ocean depths. There was little basis for this

statement, and he was proven wrong when telegraph cables were retrieved from depths

exceeding 1.7 km deep, with unknown life-forms growing on them. In 1877, Alexander Agassiz

collected and catalogued marine animals as deep as 4,240 m. He studied their coloration patterns

and theorized the absorption of different wavelengths at depth. He also noted similarities

between deepwater organisms on the east and west coast of Central America and suggested that

the Pacific and Caribbean were once connected.

Modern marine science is generally considered to have started with the HMS Challenger

expedition, led by the British Admiralty between 1872 and 1876. During a circumnavigation that

lasted 3.5 years, the Challenger sailed on the world’s oceans, taking samples in various

locations. The information collected was enough to fill 50 volumes that took 20 years to write.

The samples taken during the Challenger expedition led to the identification of over 4,700 new

species, many from great depths, and the chief scientist, Charles Wyville Thomson, collected

plankton samples for the first time.

BIOL 181: Life in the Oceans – Lecture Notes

The Challenger expedition was the start of modern marine biology and oceanography, and is still

to date the longest oceanography expedition ever undertaken. However, modern technology has

allowed us to sample organisms more easily and more effectively and to quantify things more

accurately. Scuba diving and submersibles are used to directly observe and sample marine life

remote sampling can be done with nets, bottles, and grabs from research vessels, and satellites

are used extensively for remote sensing.

1.3. Why Study Marine Biology?

1.3.1. To Dispel Misunderstandings about Marine Life

Though many people fear sharks, in reality 80 percent of shark species grow to less than 1.6 m

and are unable to hurt humans. Only three species have been identified repeatedly in attacks

(great white, tiger and bull sharks). There are typically only about eight to 12 shark attack

fatalities every year, which is far less than the number of people killed each year by elephants,

bees, crocodiles, or lightning.

1.3.2. To Preserve Our Fisheries and Food Source

Fish supply the greatest percentage of the world’s protein consumed by humans, yet about 70

percent of the world’s fisheries are currently overfished and not harvested in a sustainable way.

Fisheries biologists work to estimate a maximum sustainable yield, the theoretical maximum

quantity of fish that can be continuously harvested each year from a stock under existing

(average) environmental conditions, without significantly interfering with the regeneration of

fishing stocks (i.e., fishing sustainably).

1.3.3. To Conserve Marine Biodiversity

Life began in the sea (roughly 3-3.5 billion years ago), and about 80 percent of life on earth is

found in the oceans. A mouthful of seawater may contain millions of bacterial cells, hundreds of

thousands of phytoplankton, and tens of thousands of zooplankton. The Great Barrier Reef alone

is made of 400 species of coral and supports over 2,000 species of fish and thousands of

1.3.4. To Conserve the Marine Environment

Each year, three times as much trash is dumped into the world’s oceans as the weight of the fish

caught. There are areas in the North Pacific where plastic pellets are six times more abundant

than zooplankton. Plastic is not biodegradable and can kill organisms that ingest it. Many

industrial chemicals biomagnify up the food chain and kill top predators. Some chemicals can

bind with hormone receptors and cause sex changes or infertility in fish. Understanding these

links allow us to better regulate harmful activities.

1.3.5. To Conserve the Terrestrial Environment

BIOL 181: Life in the Oceans – Lecture Notes

Phytoplankton and algae use CO2 dissolved in seawater in the process of photosynthesis, and

together are much more important than land plants in global photosynthetic rates. Marine

photosynthesizers therefore have the ability to reduce the amount of CO2 dissolved in the oceans

and consequently in the atmosphere, which has important implications for the entire biosphere.

Many marine habitats, such as coral reefs and mangroves, also serve to directly protect coastlines

by acting as a buffer zone, reducing the impact of storm surges and tsunamis that may threaten

Because the architecture and chemistry of coral is very similar to human bone, it has been used

in bone grafting, helping bones to heal quickly and cleanly. Echinoderms and many other

invertebrates are used in research on regeneration. Chemicals found in sponges and many other

invertebrates are used to produce several pharmaceutical products. New compounds are found

regularly in marine species.

Several species of plankton are toxic and responsible for shellfish poisoning or ciguatera.

Understanding the biology of those species allows biologists to control outbreaks and reduce

their impact on human health.

1.3.8. Because Marine Organisms Are Really Cool

Many fish are hermaphrodites and can change sex during their lives. Others, including several

deep-sea species, are simultaneous hermaphrodites and have both male and female sex organs at

The blue whale is the largest animal to have ever live on earth, and has a heart the size of a

An octopus recently discovered and as of yet unnamed has the ability to mimic the color and

behavior of sole fish, lionfish, and sea snakes, all toxic animals, which greatly reduces its

likelihood of encountering predators.

1.4. How Is Marine Biology Studied? Using the Scientific Method

The word science comes from the Latin (scientia) and means ―knowledge.‖ Science is a

systematic enterprise that builds and organizes knowledge in the form of testable explanations

and predictions about the world.

1.4.2. The Scientific Method

BIOL 181: Life in the Oceans – Lecture Notes

The scientific method is widely used in the process of conducting science. Its general steps are to

make observations, form a hypothesis to explain the patterns seen, perform experiments to test

the hypothesis, and then draw conclusions (Figure 1.1).

Diagram showing the steps of the scientific method

by Erik Ong is licensed under CC BY-SA 3.0

Figure 1.1. Scientific method. Steps of the scientific method.

1.5. Review Questions: Introduction to Marine Biology
BIOL 181: Life in the Oceans – Lecture Notes

1. What percentage of the earth is covered with oceans? 2. What was the driving force behind the initial studies into oceanography? 3. Who was the scientist on board the HMS Beagle in 1831? 4. What theories did this scientist develop? 5. In the early nineteenth century, who proposed that no life could live in the deep ocean? 6. Who was the chief scientist on board the HMS Challenger from 1872 to 1876? 7. What theories did Alexander Agassiz develop? 8. Why study marine biology? Give three reasons. 9. Explain the process of the scientific method.

BIOL 181: Life in the Oceans – Lecture Notes

2. Fundamentals of Ecology (The majority of the text below originally appeared as chapter 2 of Introduction to Marine Biology)

Ecology (from Greek oikos meaning home) is the study of interactions of organisms with each

other and with their environment.

Ecosystems are composed of living organisms and their nonliving environment, while the

biosphere includes all of the earth’s ecosystems taken together.

The environment is all the external factors that act on an organism:

 physical (abiotic): temperature, salinity, pH, sunlight, currents, wave action, and sediment

 biological (biotic): other living organisms and their interactions, e.g., competition and reproduction

The habitat is the specific place in the environment where the organism lives e.g., rocky or

sandy shore, mangrove, coral reefs. Different habitats have different chemical and physical

properties that dictate which organisms can live there.

Niche: what an organism does in its environment—range of environmental and biological factors

that affect its ability to survive and reproduce

 physical: force of waves, temperature, salinity, moisture (intertidal)

 biological: predator/prey relationships, parasitism, competition, organisms as shelter

 behavioral: feeding time, mating, social behavior, young bearing

2.2. Environmental Factors that Affect the Distribution of Marine Organisms

2.2.1. Maintaining Homeostasis

All organisms need to maintain a stable internal environment, even though their external

environment may be changing continuously. Factors such as internal temperature, salinity, waste

products, and water content all need to be regulated within a relatively narrow range if the

organism is to survive. This regulation of the internal environment by an organism is termed

homeostasis. The ability to maintain homeostasis limits the environments where an organism can

survive and reproduce. Each species has an optimal range of each environmental factor that

affects it. Outside of this optimum, zones of stress exist where the organism may fail to

reproduce. At even more extremes lie zones of intolerance, where the environment is too extreme

for the organism to survive at all.

BIOL 181: Life in the Oceans – Lecture Notes

Sunlight plays an essential role in the marine environment. Photosynthetic organisms are the

base of nearly every food web in the ocean and are dependent upon sunlight to provide energy to

produce organic molecules. Light is also necessary for vision, as many organisms rely on this to

capture prey, avoid predation, and communicate, and for species recognition in reproduction.

Excessive sunlight can, however, be detrimental to some life forms, as it may increase

desiccation in intertidal areas and induce photo-inhibition through pigment damage to

photosynthetic organisms in the very top of the water column.

Click the link to see a graph showing the ranges of environmental “comfort zone.” Where planets

are concerned, the central location is referred to as the “Goldilocks Zone”: not too hot, not too

cold range of conditions for organisms.

Most marine organisms are ectotherms (meaning that they rely on environmental heat sources)

and as such are increasingly active in warmer temperatures. Marine mammals and birds, on the

other hand, are endotherms and obtain heat from their metabolism. To keep this heat, they often

have anatomical adaptations such as insulation. The temperature of shallow subtidal and

intertidal areas may be constantly changing, and organisms living in these environments need to

be able to adapt to these changes. Conversely, in the open oceans and deep seas, the temperatures

may remain relatively constant, so organisms do not need to be as adaptable.

Salinity is the measure of the concentration of dissolved organic salts in the water column and is

measured in parts per thousand (‰). Organisms must maintain a proper balance of water and

salts within their tissue. Semipermeable membranes allow water but not solutes to move across

in a process called osmosis. If too much water is lost from body cells, organisms become

dehydrated and may die. Some organisms cannot regulate their internal salt balance and will

have the same salinity as their external environment these are termed osmoconformers. These

organisms are most common in the open ocean, which has a relatively stable salinity. In coastal

areas where the salinity may change considerably, osmoregulators are more common.

At sea level, pressure is 1 atm. Water is much denser than air, and for every 10 meters descent

below sea level, the pressure increases by 1 atm. Thus, the pressure at 4,000 m will be 401 atm,

and in the deepest part of the oceans at nearly 11,000 m, the pressure will be about 1,101 atm.

The pressure of the water may affect organisms that both live in or visit these depths. Organisms

found in the deep oceans require adaptations to allow them to survive at great pressures.

Organisms need a variety of organic and inorganic materials to metabolize, grow, and reproduce.

The chemical composition of saltwater provides several of the nutrients required by marine

organisms. Nitrogen and phosphorous are required by all photosynthesizing plants or plant-like
BIOL 181: Life in the Oceans – Lecture Notes

organisms. Other minerals such as calcium are essential for the synthesis of mollusk shells and

coral skeletons. Although nutrients are essential for life, excessively high levels of nutrients in

sea water can cause eutrophication. This process of nutrient enrichment can lead to vast algal

blooms that eventually die and start to decompose. The decomposition may deplete the available

dissolved oxygen in the water, killing fish and other organisms.

2.3. Populations and Ecology

Population: a group of organisms of the same species that occupies a specific area. Different

populations are separated from each other by barriers that prevent organisms from breeding.

Biological community: populations of different species that occupy one habitat at the same

time. The species that make up a community are linked in some way through competition,

predator/prey relationships and symbiosis.

2.3.1. Population Range and Size

Since biologists can’t count every single individual in a population, they must instead estimate

size by sampling. One common way to sample a population is to count all individuals within a

few representative areas, and then extrapolate to the total number of individuals that are likely to

be in the entire range. Of course, this method only works well if the samples are representative of

the overall density of the population if you happen to sample areas of exceptionally high

density, you would overestimate population size. Another common method to estimate

population size is the mark-recapture method. In this process, a certain number of individuals are

captured and tagged, then released back and allowed to mingle with the rest of the population.

After a certain period, a second sample is taken. As long as the marked individuals are dispersed

well within the population and haven’t suffered mortality from the first capture, the ratio of

marked: unmarked individuals in the second capture should reflect the ratio of marked: unmarked

individuals in the entire population. (Click the link to see a graphic demonstrating a simple mark

and recapture model for determining population density and distribution.) Therefore, we can

estimate population size with the following formula:

M = Number of animals captured and marked in first sample

R = Number of animals captured in resampling event

m = Number of “R” that were already marked

2.3.2. Distribution of Organisms in a Population

Population density refers to the number of individuals per unit area or volume. In many

populations, individuals are not distributed evenly, and the dispersion (pattern of spacing among
BIOL 181: Life in the Oceans – Lecture Notes

individuals) can tell a lot about the spacing of resources and interactions between individuals. A

clumped dispersion pattern may reflect variations in the physical environment, or a clumped food

source a uniform distribution is often the result of strong intraspecific competition random

dispersion reflects weak interactions among individuals.

2.3.3. Changes in Population Size

Populations change in size over time. They acquire new individuals through immigration and

births, and lose individuals through emigration and deaths. Different species can have varying

reproductive outputs, life span and generation times, all of which can affect how quickly

populations of that species can grow. Collectively, these traits and others that impact births,

deaths and reproduction are referred to as life history traits. On each extreme of a continuum of

life history, strategies are r-selected species (those that have short generation times, high

reproductive potential) and K-selected species (those that have much longer generation time and

are long-lived, but have low reproductive outputs and low population growth potential. The

typical traits of r- and K-selected species are outlined in table 2.1.

Table 2.1. Characteristics of Organisms that Are Extreme r or K Strategists

Unstable Environment Density-

Stable Environment Density-Dependent

organism is small organism is large

energy used to make each individual is low energy used to make each individual is high

many offspring are produced few offspring are produced

organisms have early maturity organisms have late maturity, often after a

prolonged period of parental care

organisms have a short life expectancy organisms have a long life expectancy

each individual reproduces only once individuals can reproduce more than once in a

organisms have a type III survivorship pattern,

in which most die within a short time, but a

organisms have a type I or II survivorship

pattern, in which most live to near the

Source: Adapted from University of Miami Department of Biology. Accessed July 15, 2014, from

BIOL 181: Life in the Oceans – Lecture Notes

Populations change in size due to births, deaths, immigration, and emigration. Click here for a

graphic showing various factors that influence population size.

There are many ways in which a population can increase in size, including reproduction and

immigration. When a population has sufficient food or nutrients and is not greatly affected by

predation, it can grow rapidly in an exponential curve. However, no population can maintain this

growth forever—at some point resources become limited and slow down population growth

(either through lower birth rate or increased death rate). That population growth model is called

logistic growth. Here, the population levels off at size which the environment can sustain, known

as the carrying capacity of the environment. The carrying capacity is a dynamic point which may

fluctuate with changes in resource availability and predator behavior. Predator abundance often

mirrors prey abundance with somewhat of a lag in time. Click here to see a graph of exponential

(geometric) and logistic growth curves.

A biological community comprises the various populations of different species that interact

together in the same place at the same time. Organisms in a community interact with one another

The niche of an organism is often described as its role in the community. It refers to the

environmental conditions and resources that define the requirements of an organism. The

broadest niche that an organism can occupy (defined mostly by resource availability and

tolerance to abiotic factors, e.g., pH, salinity) is called its fundamental niche. In reality,

organisms often occupy a smaller subset of their fundamental niche because of biological

interactions with other species such as competition and predation. This subset is called the

realized niche. Click the link to see a graphic indicating the difference between realized and

fundamental niches in nature and how these zones are determined.

2.4.2. Biological Interactions

Competition—occurs when organisms require the same limiting resources such as food, space

or mates. Interspecific competition occurs between organisms of different species, whereas

intraspecific competition is between organisms of the same species. Interspecific competition for

resources prevents different species from occupying exactly the same niche if two species have

the same requirements, one will outcompete the other with several possible results: local

extinction (also known as competitive exclusion), displacement of the less successful competitor,

or selection for speciation that would lessen the competition.

To efficiently take advantage of a common resource, organisms may have unique anatomical and

behavioral specializations. This is commonly seen on coral reefs. For example, fairy basslets,

brown chromis, and soldierfish are all plankton feeders, but they do not directly compete with
BIOL 181: Life in the Oceans – Lecture Notes

each other. Fairy basslets feed close to the reef, chromic feed in the water column, and

soldierfish feed mainly at night. Another strategy to lessen competition is to take advantage of a

resource not in demand by other species, e.g., angelfish are one of the only reef fish that eat

Predator-prey relationship—may determine the abundance of different trophic levels. The

amount of vegetation in a given area may determine the number of herbivores, which in turn may

limit or be limited by the amount of primary consumers and so on and so forth up the trophic

ladder. In some situations, if the population of primary consumers becomes large and consumes

many of the herbivores, then the vegetation of the area may thrive in a process known as a

trophic cascade. This would be an example of a top-down process in which the abundance of

prey taxa is dependent upon the actions of consumers from higher trophic levels. Bottom-up

processes are functioning when the abundance or diversity of members of higher trophic levels is

dependent upon the availability or quality of resources from lower levels. For example, the

amount of algae produced determines the amount of herbivorous fish produced, and this in turn

determines the amount of piscivorous fish the ecosystem will support.

A keystone species is an organism whose effect on the biological diversity of an area is

disproportionate to its abundance. For instance, the ochre sea star (Pisaster ochraceus) in the

intertidal zone of western North America is a keystone predator and makes it possible for many

other organisms to live through its predation on the mussel. Without the ochre sea star, the

intertidal zone becomes dominated by mussels, which outcompete most other species.

Symbiosis—occurs where organisms develop close relationships to each other, to the extent that

one frequently depends on the other for survival. There are three types of symbioses:

(a) Mutualism: both organisms benefit from the relationship e.g., corals and zooxanthellae clown fish and sea anemones

(b) Commensalism: one organism benefits while the other is not harmed but doesn’t benefit e.g., remoras and sharks

(c) Parasitism: parasites live off a host, which is harmed e.g., worms in digestive tract

Ecosystems include the biological communities and their physical environment. Examples of

ecosystems include coral reefs, mangroves, rocky shores, sandy beaches, estuaries, kelp forests,

or the open ocean. Since different ecosystems don’t exist in complete isolation from one another,

important interactions between different ecosystems often exist (e.g., many coral reef fish spend

their juvenile stages in nearby mangroves).

Most producers obtain their energy from the sun or some form of chemicals. The vast majority of

primary producers photosynthesizes using a pigment called chlorophyll, which absorbs the sun’s

energy and convert it into an organic molecule called glucose (C6H12O6). Other autotrophs may

be chemosynthetic, using the energy from chemical reactions to produce organic compounds.

BIOL 181: Life in the Oceans – Lecture Notes

The glucose produced by autotrophs may be used by the organism for its own metabolic needs or

is available for higher trophic levels.

2.5.2. Consumers (Heterotrophs)

Organisms that rely on other organisms for food are collectively known as heterotrophs. Primary

consumers are herbivores, feeding on plants. Secondary consumers are carnivores feeding on the

herbivores. Tertiary consumers then feed on the secondary consumers and so on until the top

carnivores are at the top of the food chain. There are also omnivores, which feed on both

producers and heterotrophs, and then decomposers, which feed on all organic matter, breaking it

back down to simple molecules.

Click here for a graphic showing the difference between a terrestrial food chain and a marine

2.5.3. Food Chains and Food Webs

Food chains are simple representations of the feeding relationships in an ecosystem. They show

one organism feeding on one prey while being devoured by one predator. In reality, these

interactions may be much more complex with one organism feeding on several prey at different

trophic levels while having several potential predators. This more complex relationship is called

2.5.4. Other Energy Pathways

Not all energy pathways in the marine environment involve one organism feeding on another.

Through several inefficient feeding and metabolic mechanisms, organic matter is released into

the marine environment in the form of dissolved organic matter (DOM). These energy-rich

organic molecules can be incorporated by bacteria and other small plankton, which in turn are

eaten by larger organisms. In this way DOM, which would otherwise be lost to the environment,

is funneled back into the food web. Detritus from feces and decaying plants and animals is also

an extremely important food source for organisms in the marine environment. Detritivores, such

as bacteria and zooplankton in the pelagic zone or animals in the benthos, feed on this detritus,

returning energy back into the food chains.

Energy flows from the sun through producers to higher orders of consumers. Energy received

from photosynthesis, or from food, is temporarily stored until the organism is eaten or dies and is

decomposed. Thus energy storage in an organism can be portrayed as a trophic level. Primary

producers represent the first trophic level primary consumers the second, secondary consumer

the third, and so on. Energy transfer between trophic levels is inefficient primary producers

capture and store less than 1 percent of the sun’s energy. From there, an average of only 10

percent of the energy is passed on to successive higher trophic levels while the rest is used for

feeding, metabolism, reproduction, etc.
BIOL 181: Life in the Oceans – Lecture Notes

2.6.1. Distribution of Marine Communities

Marine communities and ecosystems can be designated by the regions of the oceans that they

inhabit (click here for a simple representation of the ecological energy pyramid, showing energy

loss at different trophic levels). In the water column, known as the pelagic zone, the area of water

overlying the continental shelf is known as the neritic zone, whereas the area above deep ocean

basins is known as the oceanic zone. The organisms that inhabit the pelagic division exhibit one

of two different lifestyles. Plankton drift with the currents, whereas nekton are active swimmers

that can move against the currents. The benthic realm can be divided into the intertidal area, the

continental shelf, and the deep. Organisms in the benthic division are either epifauna, organisms

that live on the sediment, or infauna, organisms that live within the sediment.

2.7. Review Questions: Fundamentals of Ecology

1. Define the term ecology. 2. Define the term ecosystems. 3. Give an example of abiotic factors affecting marine organisms. 4. Give an example of biotic factors affecting marine organisms. 5. Define the term habitat. 6. Define the term niche. 7. Define the term homeostasis. 8. What is an ectotherm? 9. What is an endotherm? 10. What does it mean if an organism is anaerobic? 11. What is eutrophication? 12. What is the difference between interspecific and intraspecific competition? 13. What is resource partitioning, and give an example on a coral reef? 14. Define a keystone predator and give an example of one. 15. Name and describe the three types of symbiotic relationships. 16. What is osmosis? 17. What are osmoconformers? 18. In which type of symbiotic relationship does one organism benefit and the other is not

harmed in any way but does not benefit?

19. Define the term population. 20. What is the carrying capacity of a population? 21. What does the term neritic refer to? 22. What is plankton? 23. What does benthic refer to? 24. What does pelagic refer to? 25. What is the difference between epifauna and infauna? 26. What is an autotroph? 27. What is the equation for photosynthesis? 28. What is the primary energy source for autotrophs? 29. What inorganic nutrients do photosynthetic organisms require?
BIOL 181: Life in the Oceans – Lecture Notes

30. How do chemosynthetic organisms generate energy? 31. What do detritivores feed on? 32. What is the average percent of energy passed from one trophic level to another in a food

chain? What is the rest used for?

BIOL 181: Life in the Oceans – Lecture Notes

3. Marine Provinces (The majority of the text below originally appeared as chapter 3 of Introduction to Oceanography)

Why study the oceans? To understand “Life in the Oceans” (our course title), we need to

understand the ocean environments. They cover 71% of our planet (Figure 3.1), and play an

important role in regulating global climate through their interaction with the atmosphere.

Figure 3.1. Pacific Ocean from space. Here is an image you probably have not seen before. This

is the Pacific Ocean, with California in the upper-right corner and New Zealand in the lower-left

corner. Perhaps our planet should be named “Water” instead of “Earth” because oceans cover

71% of the planet. The Pacific Ocean is the largest ocean.

Oceans have been present for about 4 billion years, and are thought to be where life originated.

Moreover, the majority of the human population lives by the sea, and modern societies use

biological and mineral resources from the sea. Understanding the oceans is critical for optimal

and sustainable harvest of these resources.

BIOL 181: Life in the Oceans – Lecture Notes

Four major oceans have traditionally been recognized, with one additional ocean newly

World map of oceans (English version) by Pinpin is licensed under CC BY-SA 3.0

Figure 3.2. The world’s oceans.

The average depth of all oceans is 3.5 km. Pacific means peaceful or tranquil, but is inaccurately

named as the Pacific Ocean has numerous earthquakes and volcanoes along its edge (the Ring of

Fire). The Pacific is the oldest ocean, about 200 million years old, and the deepest, with an

average depth of 4.2 km. It is the largest (13,000 km wide) and covers 1/3 of the earth’s surface.

The Atlantic Ocean is half as old as the Pacific, and much smaller (6,600 km wide). It is 3.6 km

deep on average. The Indian Ocean is 7,000 km wide and has an average depth of 3.7 km. It is

confined to the Southern hemisphere. The Arctic Ocean is frozen but does not have any land

masses. It has an average depth of 1.1 km. Most oceanographers now also recognize the

Southern Ocean as a separate ocean. Although it is physically connected to the Pacific, Atlantic,

and Indian Oceans, this body of water, south of about 50 degrees south, is defined by the distinct

circulation of the Antarctic convergence.

Seas are bodies of salt water that are smaller than oceans. They have a direct connection to an

ocean and are often indentations into continents, or delineated by an island arc. There are many

seas around the world, including the Caribbean, Mediterranean and Red Sea

3.2. Determining Ocean Bathymetry
BIOL 181: Life in the Oceans – Lecture Notes

Bathymetry is the study of the depth and shape of the bottom of the ocean. Depth can be

measured in various ways. The earliest depth readings were done using soundings: lowering a

heavy weight on a line until it reached the bottom. In the early 1900s, the first echo sounders

measured ocean depth by sending a sound signal to the bottom and measuring how long it takes

for its echo to return to the surface. Modern echo sounders have tremendously increased the

precision of the measurements, but these measurements are severely limited by ship time and

resources. For this reason, satellite remote-sensing is increasingly used to infer bathymetry.

Features on the ocean floor create sea level abnormalities above them, which can be measured

accurately by satellites after correcting for waves, tides and other interferences (Figure 3.3).

Because satellites can obtain much more data than ships, bathymetric charts derived from

satellite data are more much detailed than those produced by acoustics alone (Figure 3.4).

The Jason-1 Measurement System by NASA

is in the public domain in the United States.

Figure 3.3. The Jason-1 measurement system.
BIOL 181: Life in the Oceans – Lecture Notes

Global seafloor topographic map by NOAA

is in the public domain in the United States.

Figure 3.4. Global bathymetric chart derived from sea surface abnormalities, which reveals

continental shelves and other shallow areas in pink, mid-ocean ridges in yellow-green and the

deepest parts of the ocean in blue.

3.3. Features of the Continental Margins

The ocean floor can generally be divided in three regions: continental margins, the ocean basins,

and mid-ocean ridges (Figure 3.5). The submerged edges of continents and the steep slopes that

lead to the sea floor, both made of continental crust, are the continental margins. Continental

margins may be passive or active. Passive margins are found where continents have rifted apart

(e.g., the Atlantic). Passive margins show little seismic or volcanic activity, and the transition

from continental to oceanic crust occurs on the same plate. They are typically wide. Active

continental margins, on the other hand, are associated with convergent plate boundaries and

subduction of oceanic crust beneath continental crust (e.g., Pacific Ocean). Active continental

margins are associated with earthquakes and volcanoes, and are typically narrow.
BIOL 181: Life in the Oceans – Lecture Notes

Diagram Representing Oceanic Basin by Chris_Huh

is in the public domain in the United States.

Figure 3.5. Oceanic basin features.

Continental margins are made up of several sections (Figure 3.6). The continental shelf lies right

at the edge of the continent and is nearly flat, with an average depth of 130 m. The width of the

continental shelf varies greatly, and is much greater in passive continental margins. Continental

shelves have been alternately submerged and uncovered through fluctuations in sea level during

glacial ages, and when inundated, they may accumulate sediment derived from land and carried

by rivers. The shelf break marks the abrupt change in slope from the nearly flat continental shelf

to the continental slope. The angle of the slope varies greatly. Continental slopes have submarine

canyons that were formed during periods of low sea level (Figure 3.7). These canyons are V-

shaped with steep walls and transport sediments from the shelves to the deep sea floor. Caused

by earthquakes or overloading of sediments on the shelf, turbidity currents are a fast moving

flows of sediments on the continental slope that may travel to speeds of 90 km/hr and carry

enormous quantities of sediments. At the base of the continental slope the accumulation of

sediment creates a gentle slope. This portion of the continental margin is known as the

continental rise, and is most prominent on passive continental margins. The continental rise

marks the beginning of true deep ocean basins.
BIOL 181: Life in the Oceans – Lecture Notes

Continental Shelf by the Office of Naval Research,

U.S. Department of the Navy, is in the public domain in the United States.

Figure 3.6. Continental margin, with continental shelf, slope, and rise.

Los Angeles Margin: Perspective View Looking North

by USGS is in the public domain in the United States.

Figure 3.7. Perspective view looking north over the San Gabriel (A) and Newport (B) submarine

canyons. The distance across the bottom of the image is about 17 km with a vertical exaggeration

of 6x. Both canyons formed when the San Gabriel River and the Santa Ana River flowed out

across the Los Angeles Basin and offshore shelf when it was exposed during lower eustatic sea
BIOL 181: Life in the Oceans – Lecture Notes

level. Newport Canyon begins less than 360 m from shore at the north end of Newport Harbor

and is composed of individual channels that braid down the slope over a width of about 9 km.

San Gabriel Canyon begins as a series of channels that join together midway down the slope and

then split into two channels at the base of the slope. The width of San Gabriel Canyon at “C” is

815 m and incises about 25 m into the slope. Lasuen Knoll can be seen in the foreground.

3.4. Features of Deep Ocean Basins

The deep sea floor covers a huge area of the oceans, and most of it consists of vast flat plains

known as the abyssal plains (Figure 3.8). Sediments carried from continental shelves are

eventually deposited on the deep sea floor, covering irregular topography and forming this flat

abyssal plain. Abyssal hills and seamounts are scattered throughout the sea floor. Abyssal hills

are short (less than 1,000 m high) and are a very common feature of the deep oceans. Most are

volcanic in origin. Seamounts are steep volcanoes that sometimes pierce the surface of the water

and become islands. Submerged seamounts that have a flat top are known as guyots, and were

formed by wave erosion when they were at the surface (Figure 3.8). Deep sea trenches occur

along convergent plate boundaries and typically have steep sides. The deepest part of the oceans,

known as the Challenger Deep, occurs in the Mariana trench off Japan and is 11,020 m deep.

Ocean trenches are associated with volcanic arcs, on the side of the overriding plate, as material

from the subducted plate melts and rises. These volcanoes can form island arcs such as the

eastern Caribbean or volcanic mountain ranges such as the Andes. The Pacific Ocean is lined

with such trenches, which create volcanoes and earthquakes around its perimeter, which has been

dubbed the Pacific Ring of Fire (Figure 3.9).

Image courtesy of Prof. Denny Whitford, UMUC.

Figure 3.8. Ocean floor features. Abyssal plains comprise about 40% of the total area in the

BIOL 181: Life in the Oceans – Lecture Notes

Pacific Ring of Fire by Gringer is in

the public domain in the United States.

Figure 3.9. Deep sea trenches are especially common around the perimeter of the Pacific Ocean,

but are also found in the Indian and Atlantic Oceans. The perimeter of the Pacific Ocean is

known as “The Ring of Fire” because of earthquake activity resulting from tectonic plate

3.5. Features of the Mid-Ocean Ridges

Mid-ocean ridges and rises are the longest continuous mountain chain on earth and are

approximately 75,000 km long. Mid-ocean ridges are typically 2–3 km high and have a central

rift valley along the axis of spreading (Figure 3.10). A prominent feature of the rift valley is

hydrothermal vents (Figure 3.11a and b). These unique features are created when water seeps

down in cracks in the crust, gains heat and dissolved substances and is released by through the

seafloor. Hydrothermal vents can reach temperatures of over 350 °C and contain energy-rich

inorganic compounds such as hydrogen sulfides which can be used as a source of energy by

specialized communities than inhabit the vents. Volcanic seamounts can also be associated with

mid-ocean ridges, as magma can escape through the oceanic crust by side chambers (Figure
BIOL 181: Life in the Oceans – Lecture Notes

Age of the Sea Floor with Shaded Vegetation by

NOAA is in the public domain in the United States.

Figure 3.10. Sea floor bathymetric features and tectonic plate names. Mid-ocean ridges are

shown in red with a black line. Red indicates youngest sea floor age.

New Model for Water Dynamics of Deep-Sea Hydrothermal Vents

by Zena Deretsky, NSF, is in the public domain in the United States.

Black Smoker at a Mid-Ocean

Ridge Hydrothermal Vent by P.

Rona, NOAA, is in the public

domain in the United States.

Figure 3.11a and b. Structure and image of a hydrothermal vent.
BIOL 181: Life in the Oceans – Lecture Notes

A Map of Seamount in the Arctic Ocean by

NOAA/NOS is licensed under CC BY 2.0

Figure 3.12. Seamounts rising from the seafloor. Color code indicates depth, with red and yellow

being the shallowest depth, and purple representing the deepest depths.

Mid-ocean ridges are cut by a number of fracture zones, parallel series of linear valleys

perpendicular to the ridge. Transform faults are the region of the fracture zone where plates

move in opposite direction (Figure 3.13). Earthquakes are frequent along transform faults.
BIOL 181: Life in the Oceans – Lecture Notes

Tectonic Plate Boundaries by Jose F. Vigil (USGS, Smithsonian Institution,

US Naval Research Laboratory) is in the public domain in the United States.

Figure 3.13. Mid-ocean ridge (or oceanic spreading ridge) with multiple plate boundary types.

Note the transform plate boundaries, resulting in fracture zones (or transform faults) seen as lines

perpendicular to the ridge axis.

1. Which type of continental margin (passive or active) typically has a wide continental shelf?

2. What does bathymetry mean? 3. Which ocean has passive margins? 4. Which ocean has a lot of volcanoes and earthquakes along its margins? 5. What is the approximate average depth of a continental margin? 6. What is the name of the boundary between the continental shelf and the continental

7. What are abyssal plains very flat? 8. Where are hydrothermal vents typically located? 9. Where are rift valleys located? 10. What is the difference between a transform fault and a fracture zone? 11. What size of sediments are typically found on the deep sea floor (fine or coarse)? 12. What is the continental rise? 13. What is a turbidity current?
BIOL 181: Life in the Oceans – Lecture Notes

4. Seawater (The majority of the text below originally appeared as chapter 6 of Introduction to Oceanography)

4.1. Properties of Seawater

Salinity refers to the amount of inorganic material dissolved in water. This excludes sediments

held in suspension since those particles are not dissolved. The polar nature of the water molecule

allows it to readily dissolve salts. As salts (e.g., NaCl) are added to water, they dissociate into

and Cl ˗ ), and bond with water molecules. Water can hold a certain quantity of salt

in solution this is called the saturation value. An increase in temperature increases the saturation

On average the ocean has a salinity of 35 ‰, (ppt, parts per thousand), which means that 1,000 g

of seawater is composed of 965 g of water and 35 g of dissolved solids (Figure 4.1). The most

abundant salts in water are referred to as major ions. Note that the six most abundant of the

major ions make up 99% of all salts in seawater. Minor constituents include more ions, as well as

some gases and nutrients. Trace elements are present in concentrations lower than 1 ppm, and

include aluminum, copper, cobalt, iron, mercury and silver, among others. Though trace

elements are present only in very small quantities, they may still play an important role

Proportion of salt to sea water and chemical composition of sea salt

by Hannes Grobe is licensed under CC BY-SA 2.5
BIOL 181: Life in the Oceans – Lecture Notes

Figure 4.1. Major ions in seawater. Note that ‰ means “parts per thousand” and % indicates

“parts per hundred.” Therefore 3.5% equals 35 ‰.

The principle of constant proportions states that the ratio of one major ion to another remains the

same, regardless of variations in salinity. This applies to major conservative ions in open-ocean,

not where rivers bring dissolved substances or reduce salinities. The ratios of minor non-

conservative constituents (e.g., nutrients and gases) do not follow the principle of constant

composition as they vary because they are connected to life cycles of organisms.

The salinity of water can be measured in a variety of ways. It can be measured with a

salinometer, which measures the conductivity of water (the saltier the water, the more it conducts

electricity). CTDs (Conductivity-Temperature Devices) are modern instruments that also

measure salinity through conductivity. It is also possible to titrate the chlorine in the water,

which is directly proportional to the total salinity because of the principle of constant

proportions. A refractometer measures the bending of light as it passes from air to water the

saltier the water, the more dense it is, and the more it refracts light.

Pure Water versus Seawater

Seawater has slightly different properties than pure water: it freezes at a lower temperature, boils

at a higher temperature, has higher density and higher pH.

4.2. Processes Affecting Salinity

In the open ocean, salinity varies slightly, from about 33 to 38‰. Salinity variations are much

more extreme in coastal areas, where freshwater input and evaporation can create brackish water

(between seawater and freshwater) or hypersaline water (e.g., up to about 42‰ in the Red Sea).

Salinity often varies seasonally based several factors that affect water input or removal (and to a

much lesser degree, salt input or removal).

Salts present in the oceans originally came from the crust and interior of the earth and were

released through volcanism and hydrothermal vents. Physical and chemical weathering of rocks

on land also adds salt to the oceans through river runoff.

It is estimated that the oceans have been present for about 3.5 billion years, and it appears that

salinity has remained stable for the last 1.5 billion years. If salts are continuously added through

the processes explained above, then they must also be removed by other processes for salinity to

be stable (See an illustration of these processes). Removal of salts occurs in various ways. Sea

spray leaves some salt on land, which is removed from water. Shallow seas that evaporate over a

long period of time leave salt deposits (evaporites). Biological organisms concentrate ions in

their feces and shells, which may then be transferred to sediments. Salts in sediments may be

returned to the interior of the earth as tectonic plates collide and one plate is subducted (pulled)

under the other, along with some of the sediment overlying it. Finally, ions can adhere on the
BIOL 181: Life in the Oceans – Lecture Notes

surface of small particles, e.g., clay, in a process called adsorption. Salts are then incorporated in

sediments and do not return to seawater readily.

The residence time is the average time a substance remains in solution in the ocean. Conservative

constituents have long residence times (e.g., millions of years), because they tend to be non-

reactive with water and are not added or removed by biological processes. Conservative

constituents include major ions as well as some trace elements and some gases. Non-

conservative constituents, on the other hand, are typically tied to biological, seasonal or

geological cycles. They have a short residence time ( 25 m depth or fry and spat using cast nets. No anchoring of

 Replenishment Zones (5,214 ha)—Line fishing and anchoring permitted so long as chain does not touch coral. No taking of conch or lobster.

 Designated Grouper Spawning Sites—No fishing of grouper during winter spawning season (November 1—March 31).

 Animal Sanctuaries (on land)—No hunting, no collection of any species, and no littering.

BIOL 181: Life in the Oceans – Lecture Notes

Dense Cover of Brittle Stars, by Peter Southwood, is available under a

Creative Commons Attribution-ShareAlike 3.0 Unported license.

Figure 23.1. Marine protected area.

Other regulations have also been enforced regarding species size limit, maximum take and closed

seasons. This includes Caribbean lobster (Panulirus argus), which has a closed season between

February 1 to July 31, and only three individuals larger than 6 inches can be taken per person.

In order to police Cayman waters, special law enforcement officers have been created. The Royal

Cayman Islands Police have two Marine Policy Officers while the Department of Environment

has four Marine Enforcement Officers. The penalties for breaking these regulations are harsh.

Violation of any Marine Conservation Law is subject to a fine of CI$ 5,000 (US$ 6,250), a

possibility of one year in jail, and the seizure of your boat and equipment.

In the past, many MPAs have been created by consumer groups, e.g., villagers, in a bottom-up

management style. While these areas may have strong control and regulation from the

population, they often lack strong legal and financial backing. On the other scale, governments

sometimes try to impose MPAs on populations in a top-down management type. While these

sometimes will have financial and legal backing, they often fail in their support by the local

consumer groups, with poaching and illegal activities occurring. What has become clear over

time is that a co-management of top-down/bottom-up approach is best with consumer group

support and regulation backed up by governmental legal and financial backing. Once established,

MPAs have the potential to finance themselves through park fees and user permits.

A considerable number of measures are available for the protection of coral reefs, including

fisheries controls, protected areas, and other schemes ranging from diver cleanups to consumer-

or market- driven controls on reef utilization. All of these measures are heavily dependent on

awareness, education, and the establishment of training programs.

Education needs to be aimed at all levels, including politicians and senior managers, artesian and

commercial fishers, recreational users of reefs, tourists, and aquarium hobbyists, but also the vast

number of people whose lifestyles and businesses may affect reefs through pollution or

The concept of integrated coastal zone management (ICZM) has been widely accepted and

promoted in many countries. In essence, this involves developing a policy, not for particular

locations but for the entire coastal zone, including inland watersheds, and also offshore waters.

Such policies, if developed with active participation of local stakeholders, can be a highly

effective means. The development of such integrated measures is critical, but also challenging,

requiring considerable coordination of different groups of people and complex negotiations and

processes of conflict resolution. Provision of further information and research remain a further

priority. This includes establishing or expanding systems to monitor coral reefs around the world

in order to establish a clearer baseline and provide warning of change.
BIOL 181: Life in the Oceans – Lecture Notes

23.7. Review Questions: Marine Protected Areas

1. Name three important values of the marine environment.

2. Name three natural threats to the marine environment.

3. How can MPAs help protect a reef from natural threats?

4. Give three anthropogenic threats to the reef.

5. What are the two broad main reasons for these anthropogenic threats?

6. What is the “tragedy of the commons”?

7. What are four benefits of MPA?

8. What is the Spill Over effect?

10. What are three possible aims of an MPA?

11. What are three questions to consider when planning an MPA?

12. List four potential stakeholders in an MPA.

13. Give three reasons why MPAs often fail.

14. Explain an effective way of financially supporting an MPA, and give an example of

15. What are some traditional management systems, and why are they beginning to fail?

16. Why is the Great Barrier Reef MPA so effective?

17. Give an example of three fishing regulations in the Cayman Islands.

18. Why is it important that the management of MPAs takes a holistic approach using

integrated coastal zone management (ICZM)?

19. What is the difference between a top-down and a bottom-up approach to marine

management? And what is the most effective solution?

20. What is meant by the term ―shifting baselines‖?

BIOL 181: Life in the Oceans – Lecture Notes

24. Impact of Tourism on the Marine Environment (The majority of the text below originally appeared as chapter 21 of Introduction to Marine Biology)

Tourism is the fastest growing industry worldwide. As more people travel around, their activities

directly and indirectly affect the environment. In this chapter, we explore how activities such as

scuba diving and boating negatively affect the marine environment.

24.1.1. History and Background

Scuba diving was developed by Jacques Cousteau and his team in the 1940s. It became a more

widely practiced sport in the 1970s with important technological advances (which included

nonreserve K-valves, single-hose regulators and BCDs) and the implementation of a certification

system. The scuba diving industry has seen incredible growth in the last few decades, and there

are now around 1 million new certified divers each year. People dive all around the world in

many different types of habitats, including lakes, quarries, and kelp forests. However, it is coral

reefs, with their impressive diversity of life and color, which are among the most dived

environments. Coral reef tourism generates revenues of $2 million/year in Saba and $682

million/year in Australia. This section will focus on the impact of divers on coral reef

24.1.2. Pressure from Divers

The density of divers is astounding in some parts of the world. In heavily used dive sites of Eilat

(Red Sea), there are over 30,000 dives per site per year, which averages to over 82 divers per

day. Studies in this area have shown that on average, each diver causes one coral break per dive.

The Cayman Islands, in the Caribbean, see 350,000 divers who collectively conduct millions of

dives each year. The Great Barrier Reef receives over 2 million visitors annually, and 15 percent

of the divers are observed damaging coral. These numbers are constantly increasing with the

increase in the number of certified divers.

24.1.3. How Divers Can Affect the Coral

Most damage caused by divers is by direct breakage caused by their fins. Touching live tissue of

the coral without breaking the skeleton is also damaging: it induces stress which requires energy

to repair, and damaged corals may be more susceptible to pathogens, so they thus have a higher

mortality. Moreover, poor buoyancy and swimming too close to the bottom can raise sediment

clouds that can later settle and smother corals. The coral animals then need to divert energy from

growth and reproduction to get rid of this sediment.

Dive tourism also means that more boats come to the reefs, many of which can cause damage to

corals through anchoring where no moorings are available. The overall impact from divers

increases the stress on the coral animals and may reduce their ability to cope with hurricanes,

storms, disease and other natural threats.

BIOL 181: Life in the Oceans – Lecture Notes

24.1.3. Actual Impact of Divers on Reefs

In Eilat (Red Sea), the proportion of broken and/or abraded coral can be up to 100 percent

around heavily used sites. In the Cayman Islands, increased number of divers has been correlated

with decreased coral cover and increased dead coral and rubble. In Bonaire, heavily dived sites

showed a higher abundance of branched corals, which tend to have a higher growth rate and are

usually the first to grow back after disturbances.

24.1.4. Which Divers Are Most Damaging to Reefs?

Student (uncertified) divers or certified divers with poor buoyancy skills are more likely to

inadvertently touch the bottom. All divers are more likely to touch the bottom in the first 10

minutes of a dive, while they are adjusting their buoyancy. Moreover, male divers have been

shown to damage the reef more than female divers. Women touch the reef more with their hands

but less with their fins, resulting in less damage. Underwater photographers also touch the reef

frequently, as they spend more time closer to the reef busy handing their cameras, and therefore

can lose focus on their buoyancy. Finally, night dives result in more damage to the reef than day

24.1.5. Which Sites Are Most Sensitive?

Shore dives tend to result in much higher contact than boat dives. Branching coral is usually

more susceptible to breakage than massive (mound-shaped) coral, but it also grows faster and

therefore recovers faster after damage.

Regulating agencies could limit the number of dives per site per year by calculating a ―carrying

capacity.‖ Opinions vary on this carrying capacity, and it has been suggested to be around 5,000

dives/site/year in Bonaire and Eilat, yet as high as 10,000 to 15,000 dives/site/year in Egypt. It is

difficult to simply calculate a carrying capacity for a given site, as diver behavior is often more

important than simply the number of divers.

Some areas require that all dives be led by guides, and ideally guides should lead by example,

staying far from the reef and not touching it. Dive leaders can intervene when they observe

contact with the reef and encourage divers to stay even farther away from the reef at night. Dive

operators can increase environmental awareness with a mandatory short briefing before dives,

which can include reef biology, contacts caused by divers and the concept of protected area.

Governments could set up strict accreditation criteria for dive operators and remove the

accreditation and license to operate if they are not following the standards.

Governments can exclude uncertified or new divers from vulnerable reefs and forbid the use of

gloves divers are less likely to touch the reef if they might hurt themselves. Dive moorings

reduce anchor damage, and the provision of entry points in nonsensitive areas allows for divers

to adjust their buoyancy at the beginning of the dive.

BIOL 181: Life in the Oceans – Lecture Notes

It has even been suggested that in particularly vulnerable or damaged areas, divers should be

encouraged to dive on artificial reefs or wrecks, even on underwater theme parks. All of these

would take pressure away from the natural reefs.

Marine parks that provide much of the support for maintaining dive moorings and regulating the

diving can ask for a fee from divers, which can allow them to be self-funded.

24.1.7. Shark/Fish Feeding Dives

It is common practice in many areas to feed fish, turtles and sharks. This ensures that divers see

them, and yet it is highly debated whether this practice should continue. Feeding the organisms

(Figure 24.1) may have ecological and health impacts on the wildlife, and could increase the risk

Fish Feeding, by Philippe Bourjon, is available under a

Creative Commons Attribution-ShareAlike 3.0 Unported license.
BIOL 181: Life in the Oceans – Lecture Notes

Ecological impacts: Feeding marine organisms disrupts their natural behavior, alters their

distribution and feeding patterns, and attracts them to a certain dive site so that feeding becomes

an event associated with people and boats.

Health impacts: The fed marine life becomes dependent on foods they often cannot digest

properly, or which would not be a part of their natural diet otherwise.

Increased risk of harm to divers? Attacks and bites have been increasingly reported by those

conducting feeding activities and others in the vicinity because the marine life may mistake

diver’s actions for handouts and/or the marine life loses their natural wariness of humans

It seems clear that feeding marine life has detrimental ecological consequences, but it is highly

debated whether shark feeding increases bites and attacks on humans. There is not enough

evidence to draw a definite conclusion. However, Florida and Hawaii have both outlawed shark

feeding dives in their waters.

There are 17 million recreational boats in the United States, from canoes to yachts. This number

has increased dramatically in the last decade. Their combined activities can have a significant

impact on the marine environment.

24.2.1 Environmental Impacts of Recreational Boats

Habitat damage: Coral reefs and seagrass beds are negatively affected by grounding, propellers

and anchors. The anchor chain can also cause damage by dragging around on the substrate with

the impacted area being quite large.

Propeller contact with marine mammals: Manatees are particularly vulnerable to propeller

injuries because they are slow and spend much time at the surface. Seventy to 100 manatees are

killed every year in Florida alone, and many more are injured.

Sewage: In the United States, boats equipped with a head are required to have holding tanks and

sewage cannot be released within 3 nm from shore. However, most areas outside US waters and

outside major ports do not strictly enforce the holding tank regulations, and many heads are

dumped directly overboard. This does not have a significant impact offshore and for small boats

near shore, as long as numbers are low. However, it may be a problem where there is a big

concentration of boats with limited water circulation (e.g., Tobago Cays in the spring).

Spilled fuel and oil: Many small boats lack a fuel gauge, and simply stop filling their tanks

when it overflows. Fuel in the water affects marine organisms in many ways. It blocks light and

reduces primary production it is toxic and enters the food chain, where it bioaccumulates and

biomagnifies, becoming more concentrated at higher levels of the food chain. Eventually it kills

fish, and causes internal bleeding and death in birds and mammals.

BIOL 181: Life in the Oceans – Lecture Notes

Food waste: Though food is clearly biodegradable, high levels of food dumped in a small area

diminishes water and sediment quality, and elevates nutrient levels. If large amounts are

regularly dumped at the same place, it would possibly result in ecological changes such as

changes in species behavior and alterations to community composition. However, the input from

the typical boat is small, especially if the boat is moving in various areas this is more of a

concern for warships and cruise ships.

Products: Eighty-nine percent of the 10,500 ingredients that are used in personal care products

have not been evaluated for safety, and the impacts on marine life are unknown. Several UV

filters used in sunscreen are suspected to have endocrine disrupting effects, and products that

target hormone systems have been linked to feminization of fish and other aquatic life.

24.3. Impact of Cruise Ships

In 2003, there were 250 cruise ships around the world, collectively carrying 12 million

passengers per year. Cruises are the fastest growing sector of the tourism industry, with an 8

percent increase yearly. The Caribbean and Mediterranean are the most important destinations,

but polar regions are increasing in popularity.

In one week, a typical cruise ship generates:

 1,000,000 gallons of “gray water” from showers, sinks, dishwashers and washing machines

 37,000 gallons of oily bilge water

 More than eight tons of solid waste

 Toxic wastes from onboard operations like dry cleaners and photo processing laboratories.

Illegal discharges of oil: Though this is very strictly regulated, many cruise ships have been

found to illegally discharge their oil. The Texas Treasure is accused of having illegally

discharged waste oil and deliberately bypassed its pollution prevention equipment. Ecstasy,

Fantasy, Imagination, Paradise, Sensatio, and Tropicale were found on numerous occasions

from 1996 through 2001 to have discharged oily waste from their bilges into the sea by

improperly using pollution prevention equipment. They also falsified the oil record books in

order to conceal those practices.

Anchoring on coral reefs: In 1988, the Wind Spirit dropped anchor on a coral reef in the US

Virgin Islands, destroying an area 128 m long and 2–3 m wide.

Running aground: In 1994, the Starward discharged 100 gallons of hydraulic oil on the reef

when the ship ran aground in St. John, US Virgin Islands.

BIOL 181: Life in the Oceans – Lecture Notes

Waste: Big cruise ships generate as much waste as a small city. On average, they produce 3.5 kg

of trash per passenger per day. Many cruise ship destinations that have not signed MARPOL

(Convention for the Prevention of Marine Pollution), can refuse to accept the ship’s trash and

reduce pressure in their own landfills. Cruise ships are then more likely to dump at sea.

A large cruise ship generates 1 million liters of black water in one week. Some have treatment

facilities, but most do not. They must retain the black water until they are out to sea, but there is

some illegal dumping. The Norwegian Sun was found guilty of an illegal discharge of 16,000

gallons (40 tons) of raw sewage into the Strait of Juan de Fuca, known to be habitat for Orca

Tourism to remote destinations: Cruise ships bring tourists to remote destinations they could

not otherwise access. For example, they bring snorkelers and divers to remote reefs that would

otherwise be mostly untouched. Moreover, they bring a high number of tourists to polar

ecosystems that are quite vulnerable.

Cruise ships have been operating in Alaska and the Canadian Arctic for many years. Many are

now traveling to Antarctica, in what appears so far to be an environmentally sustainable industry.

The International Association of Antarctic Tour Operators sets up guidelines to minimize impact,

which has kept cruise ship and tourist numbers to a minimum. However, the mega cruise ship

Golden Princess (109,000 tons 3,700 passengers) is now proposed to visit Antarctica (as of

24.4. Review Questions: Impact of Marine Tourism

1. Name three ways divers can directly damage coral while diving.

2. Name three additional ways the impact of divers negatively affects the reef.

3. How does raising sediment clouds hurt corals?

4. What are five ways to reduce the impact of divers on the reefs?

5. Which types of divers are more susceptible to damage reefs?

6. Which types of dive sites or corals are more susceptible to damage?

7. What are the potential negative impacts of feeding marine organisms, e.g., sharks?

8. Explain five ways in which recreational boats negatively affect the marine environment.

9. Explain four ways in which cruise ships negatively affect the marine environment.

10. Why are oil and fuel damaging to the marine environment?

11. How is sewage potentially damaging to the marine environment?

12. Define bioaccumulation and biomagnification.

13. Which marine mammal is particularly vulnerable to propeller damage in Florida?

14. Which countries can refuse the trash from cruise ships and other boats?

15. Why should we be concerned about an influx of tourists to remote areas?

BIOL 181: Life in the Oceans – Lecture Notes

25. The Global Trade of Marine Ornamental Species (The majority of the text below originally appeared as chapter 22 of Introduction to Marine Biology)

Note: Statistical information for this chapter comes from UNEP-WCNC, From Ocean to

Aquarium: The Global Trade in Marine Ornamental Species (2003).

Between 1.5 million and 2 million people worldwide have marine aquaria, and the majority of

animals for these are collected on coral reefs. There are also aquaria in many public service

buildings such as dental offices, hospitals, restaurants, and night clubs, and there are many large

public aquaria, e.g., the Georgia Aquarium, which contains 8 million gallons of water, and more

aquatic life than any other aquarium. The aquarium trade has been heavily criticized in the past

for the use of unsustainable collection techniques and poor husbandry practices.

However, this industry provides a potential source of income for communities living close to

coral reefs and an incentive for coral reef conservation from the fishermen involved. In 2003, an

interesting report was published by UNEP World Conservation Monitoring Centre, “From Ocean

to Aquarium: The Global Trade in Marine Ornamental Species,” taking an objective look at this

25.1. When Did This Industry Begin and How Has It Developed?

The collection and export of tropical marine fish for the aquarium trade started in Sri Lanka in

the 1930s, on a very small scale. Trade expanded during the 1950s, with an increasing number of

places (e.g., Hawaii and the Philippines) issuing permits for the collection of species destined for

the marine aquarium trade.

In general, the overall value of the marine fish trade has remained fairly stable in recent years.

However, the export of live coral has increased by 12–30 percent annually from 1990 to1999,

only stabilizing since 2000.

25.2. Where Do the Fish Come from?

Over 20 million wild fish are caught every year for the aquarium trade, mainly from tropical

coral reefs in Southeast Asia, particularly Indonesia, but increasingly from several island nations

in the Indian and Pacific oceans. To date, only 1–10 percent of marine fish and less than 1

percent of coral species can be bred in captivity. Even fewer species are bred in commercial

quantities. This is a large contrast to freshwater aquaria species, where 90 percent of species are

currently farmed. Perhaps advancements in aquaculture may enable the farming of marine

ornamental species in the future.

25.3. What Organisms Are Traded?

Very few of the species traded are exploited directly for other purposes, and aquarium animals

are the highest value-added product that can be harvested from a reef.

BIOL 181: Life in the Oceans – Lecture Notes

A total of 1,471 species of fish are traded globally, and the annual trade is between 20 million–

24 million individuals. Damselfish (Pomacentridae) make up almost half of the trade, with

species of angelfish (Pomacanthidae), surgeonfish (Acanthuridae), wrasses (Labridae), gobies

(Gobiidae) and butterflyfish (Chaetodontidae) accounting for approximately another 25–30

percent. One of the most traded fish is the anemonefish (Figure 25.1), particularly after the

release of Disney’s movie Finding Nemo.

Clown Fish by Michael Johnson is available under a Creative Commons Attribution 2.0 Generic license.

Figure 25.1. Clown fish, Amphiprion ocellaris.

The bluestreak cleaner wrasse (Labroides dimidiatus) (Figure 25.2a) and the mandarin fish

(Synchiropus splendidus) (Figure 25.2b) are known to not acclimatize well to aquarium

conditions, mainly due to their restricted dietary requirements. However, according to trade data

collected for the Global Marine Aquarium Database (GMAD) (2000), they are very commonly
BIOL 181: Life in the Oceans – Lecture Notes

Cleaner Wrasse with a Client, by Alexander Vasenin, is available under a

Creative Commons Attribution-ShareAlike 3.0 Unported license.
BIOL 181: Life in the Oceans – Lecture Notes

Mandarin Fish by Steve Childs is available under a Creative Commons Attribution 2.0 Generic license.

Figure 25.2a and b. The bluestreak cleaner wrasse, Labroides dimidiatus, servicing a larger fish

(a) and two mandarin fish (Synchiropus splendidus) (b).

A total of 140 species of scleractinian corals (reef-building corals) are traded worldwide, with

the annual global trade ranging from 11 to 12 million pieces. Sixty-one species of soft coral are

also traded, amounting to close to 390,000 pieces per year.
BIOL 181: Life in the Oceans – Lecture Notes

Seashells, Coral, Shark Jaws and Dried Blowfish by Tom Oates (2008) is available

under a Creative Commons Attribution-ShareAlike 3.0 Unported license.

Figure 25.3. Coral collected for the aquarium trade.

Several species of soft corals are commonly traded due to their ability to heal wounds and

regenerate tissue rapidly. Sarcophyton spp. and Dendronephthya spp. (Figure 25.4a and b) are

two of the most commonly traded species of soft coral. The biology of Sarcophyton spp. makes it

a hardy, fast-growing and easily propagated species under aquarium conditions. Unfortunately,

Dendronephthya spp. usually die within a few weeks, mainly because they lack photosynthetic

symbionts and rely on filtering particles and nutrients in the water column for food.
BIOL 181: Life in the Oceans – Lecture Notes

Sarcophyton glaucum at Shaab Angosh Reef, by Alexander Vasenin,

is available under a Creative Commons Attribution-ShareAlike 3.0 Unported license.
BIOL 181: Life in the Oceans – Lecture Notes

Dendronephthya soft coral at Gilli Lawa Laut, by Alexander Vasenin,

is available under a Creative Commons Attribution-ShareAlike 3.0 Unported license.

Figure 25.4a and b. Sarcophyton sp (a) and Dendronephthya sp (b).

In addition to coral, a further 516 species of invertebrates are traded for aquaria. The annual

global trade ranges from 9–10 million animals, mostly mollusks, shrimps and anemones. Two

species of cleaner shrimp (Figure 25.5) and one genus of anemones accounts for 15 percent of all
BIOL 181: Life in the Oceans – Lecture Notes

Banded Coral Shrimp by Richard Ling available under a

Creative Commons Attribution-ShareAlike 2.0 Generic license.

Figure 25.5. Cleaner shrimp, Stenopus sp., one of the most commonly traded invertebrate

Linckia laevigata (Figure 25.6) is the most commonly imported sea star in the aquarium trade,

and accounts for 3 percent of the total trade in invertebrates. However, they are very difficult to

maintain in aquarium conditions, due to their dietary needs of organically enriched detritus that

typically cover live rock, and they will refuse artificial aquarium food.
BIOL 181: Life in the Oceans – Lecture Notes

Linkia Laevigata by NOAA is in the public domain in the United States.

Figure 25.6. Linckia laevigata, the most commonly traded sea star.

In order to make more informed decisions on the suitability of marine organisms for the

aquarium trade, more information is need on the population dynamics and life history

characteristics of these targeted organisms.

25.4. What Is the Global Value of the Aquarium Trade?

Although the volume of organism traded is relatively low, the value is very high. The trade of

marine ornamental species is a global multimillion dollar industry, estimated to be worth

US$200 million–$330 million annually. This can potentially provide an incentive for fishermen

to conserve reef habitats and offers a livelihood to coastal communities often in low-income

In 2000, 1 kg of aquarium fish from the Maldives was valued at almost US$500, whereas the

same weight of fish harvested for food was worth only US$6. Similarly, the live coral trade is

estimated to be worth about US$7,000 per ton, whereas the use of harvested coral for the

production of limestone yields only about US$60 per ton. In the Pacific island of Palau, live rock

is exported for the aquarium trade at US$2.2–$4.4 per kilo whereas it is sold locally as

construction material for less than US.02 per kilo. In Sri Lanka, an estimated 50,000 people

are directly involved in the export of marine ornamentals.
BIOL 181: Life in the Oceans – Lecture Notes

Live Rock, by Mrbrefast, is available under a Creative Commons Attribution-ShareAlike 3.0 Unported license.

25.5. What Are the Benefits?

The aquarium trade could potentially be a sustainable use for coral reefs, if correctly managed.

Anthropogenic impacts threaten 88 percent of all reefs, particularly in Southeast Asia, the major

source of animals for the marine ornamental trade. It is important that the collection of aquarium

species does not further compound these problems. Some collection techniques have minimal

impact on coral reefs, and well-managed shipping and husbandry practices can also minimize

The trade of marine ornamentals can also provide a valuable source of foreign exchange for

national economies and a strong economic incentive for the sustainable management of reefs.

Aquarium animals are the highest value-added product that can be harvested sustainably from

coral reefs, so collecting and exporting marine ornamentals in developing countries creates jobs

in rural, low-income, coastal areas where resources and alternative options for generating income
BIOL 181: Life in the Oceans – Lecture Notes

Domestic or public saltwater aquaria increase awareness and educate the public about coral reefs,

a mostly hidden ecosystem.

25.6. What Are the Associated Problems?

Destructive collection techniques, the introduction of alien species, possible overharvesting of

some species, high levels of mortality associated with inadequate handling and transport, the lack

of scientific information for many species collected and the threat of extinction of target species

continue to pose significant challenges to achieving sustainability.

25.6.1. Destructive Harvesting Techniques

Chemicals are often used to stun and catch fish. Sodium cyanide is squirted into crevices where

fish hide and stuns the fish, making them easier to catch. However, there are high mortality rates

post-capture due to their weakened state. Therefore, more fish are collected to allow for these

fatalities. Between 5–75 percent of fish collected using narcotics die within hours of collection,

and 20–50 percent die soon after. On average, another 30 percent die prior to export, and

importing countries register mortalities of 30 percent or more. Cyanide poisoning is nonselective

and may destroy coral reef habitat by poisoning and killing nontarget animals, including corals.

Reports have demonstrated that exposure of corals to cyanide causes bleaching.

The use of cyanide originated in Taiwan and/or the Philippines in the 1960s and in the mid-

1980s, more than 80 percent of all fish harvested in the Philippines for the aquarium trade were

collected using cyanide. Its use then spread to Indonesia, and in the mid-1990s it was estimated

that about 90 percent of vessels transporting live fish in the eastern islands of Indonesia had

cyanide on board. In the past 20 years, more than 1 million kg (1,100 tons) of sodium cyanide

has been used in reefs. This amount is enough to kill 500 million people.

Cyanide fishing is illegal in most countries. In Indonesia, for example, legislation since 1985

includes specific prohibition of the use of destructive fishing practices, such as the use of poison,

with penalties up to 10 years in prison and/or a fine equivalent to US$12,000. However, the high

premium paid, the ease with which a great number of fish can be caught in a short time period,

the often poor law enforcement capacities and high levels of corruption have allowed the use of

poison to spread rapidly throughout the Asia-Pacific region and have made the eradication of this

illegal and highly destructive fishing technique nearly impossible.

During collection of corals, many other colonies may be damaged or broken. In some cases,

coral is actually broken to allow access to fish sheltering in the reef. This tends to be more

common with branching species in which small fish, such as damselfish species, often find

Collection of live rock is potentially destructive as it may lead to increased erosion and loss of

important fisheries habitat. The results of studies of the effects of collection of live rocks on reef

habitats have been inconclusive and this is a relatively new trade, so its impacts have not been

well studied. More research and monitoring is needed.

BIOL 181: Life in the Oceans – Lecture Notes

There are also risks to the divers themselves, who often go to considerable depths for extended

periods of time and may suffer from decompression sickness, “the bends,” upon return to the

25.6.2. Overfishing of Target Organisms

There are fears that favored species of ornamental fish species have been reduced to levels that

are beyond recovery, and it is believed that collection restrictions must be implemented.

Although no marine species collected for the aquarium trade are known to have been driven to

global extinction, various studies in Sri Lanka, Kenya, the Philippines, Indonesia, and Hawaii all

reported localized depletion of a number of target aquarium species of fish (e.g., butterflyfish,

angelfish) due to heavy collecting pressure.

Corals are extremely slow growing, and mortality rates of coral pieces in home aquaria are fairly

high, although improved technology may one day increase longevity. It is important to bear in

mind that practices such as coral mining for the production of lime rock have a much more

significant impact on coral populations and community structure than the collection of corals for

Due to their distinctive bright coloration, males of many coral reef fish species tend to be

preferred as they fetch a higher price. Selectively harvesting for males on a regular basis may

lead to reproductive failure and ultimately population collapse due to heavily biased sex ratios in

25.6.3. Post-Harvesting Mortality

There are many factors that lead to post-harvesting mortality, such as physical damage and use of

chemicals during collection, poor handling practice and disease. Where organisms are collected,

stored and handled by adequately trained individuals, and transported in unsuitable conditions,

estimated levels of fish mortality have been as low as a few per cent. As a result of such

mortality, more fish must be collected to meet market demand.

The introduction of species to an area where they do not naturally occur can be a serious

problem. Species are introduced through intentional and accidental stocking, release of bait fish,

release of unwanted aquarium fish, escape from aquarium facilities and discharge of ballast

water from ships. Lionfish (Pterois volitans) (Figure 25.8) are a commonly kept aquarium

species. Their dramatic invasion of the Caribbean was due to release from an aquarium in

Florida. Lionfish are now extremely abundant in the Bahamas and some parts of the Caribbean,

and they are thought to dramatically reduce the abundance of small reef fish through predation.

BIOL 181: Life in the Oceans – Lecture Notes

Lionfish, by Altbert Kok, is in the public domain in the United States.

Figure 25.8. Lionfish (Pterois volitans). The lionfish is an invasive species in the southern

25.7. How Is the Trade of Marine Ornamentals Being Managed and How Can

Sustainability Be Improved?

Ornamental fisheries must be managed to be biologically sustainable, not conflict with other

resource uses and keep post-harvest mortalities to a minimum. In order to reach biological

sustainability, the species must replenish naturally at the same or a greater rate than they are

caught, and negative impacts on the environment must be minimized. Organisms unsuitable for

aquariums must not be collected.

Protection can be achieved through the establishment of marine reserves, in which it is illegal to

collect marine ornamentals. Other methods to control collection pressures include setting quotas

and size limits, and restricting access through the use of permits.
BIOL 181: Life in the Oceans – Lecture Notes

A number of countries (e.g., Australia, the Cook Islands, Palau) make effective use of a licensing

system to monitoring the industry and regulate collection effort by issuing a limited number of

permits each year. In Australia, there are also restrictions on net size to prevent a greater fishing

effort offsetting the limited number of permits. However, in countries such as Indonesia, permits

are issued but enforcement is difficult and so the permits are largely ineffective.

The number of fish exported from any source country can also be limited though quotas. Quotas

are relatively easy to enforce and they are only effective is based on rigorous scientific research.

They must also be implemented at a species-specific level, as general quotas may encourage

collectors to focus on the most valuable species.

Juveniles are favored in the aquarium industry due to their distinctive coloration, low transport

costs and optimal size for home aquariums. However, young fish are easily stressed and suffer

high mortality during holding and transporting. Minimal size limits help ensure stock is not

unnecessarily wasted. Maximum size limits are equally important to ensure sufficient numbers of

breeding adults remain. Maximum size restrictions for collection of coral ensures mature

colonies are not removed from the reef, and removing small colonies is less likely to damage the

habitat structure of the reef.

Marine reserves are often used to manage marine fisheries, usually food fisheries, and they have

been shown to increase fish abundance and protect ecosystems from habitat destruction.

Therefore, if managed correctly they could also be valuable in managing aquarium fisheries.

Temporary closures protect species during the reproductive phase to ensure sufficient

recruitment to sustain the population. These are only effective if implemented at the right time

and in the right location. At present, there are no closed seasons in operation specifically for the

It is important that management decisions, such as the location of reserves, involve the

participation of all stakeholders, including appropriate consultation with scientists and fishermen

at the local and national levels.

The government and industry can help support conservation initiatives however, consumers can

also encourage and promote best practices. Certification empowers consumers to assist in

reducing the environmental impacts of the trade by selectively purchasing specimens produced in

an environmentally friendly manner. The Marine Aquarium Council (MAC) is developing a

certification scheme that aims to ensure market demand and support for quality products and

sustainable practices. The scheme covers practices (industry operators, facilities, and collection

areas) and products (organisms).

Industry operators can be certified through evaluation for compliance with MAC standards for

Certification of Practices. Participating companies initially have to pay fees to an independent

certification authority as well as MAC, although this is balanced against superior returns from

certified marine ornamentals from the industry as well as the consumers’ willingness to pay a

BIOL 181: Life in the Oceans – Lecture Notes

premium for certified products. For certification systems to be effective, public awareness should

be raised as the purchasing power that hobbyists possess is undoubtedly the single most

important market force in the marine aquarium industry.

The pressure on wild populations could be relieved by supplying the industry with tank-bred

rather than wild-caught specimens.

The aquaculture of corals is achieved through fragmentation of a colony and attaching the

smaller fragments to a new substrate, which can either be left in a holding tank or placed back in

the ocean until they reach marketable size. An additional advantage is that cultured coral adapts

better to aquarium conditions than wild-caught coral. Seventy-five species of coral can be bred in

captivity, but only fast-growing species are economically profitable, e.g. Acropora and

Pocillipora. Unfortunately, most of the popular species in the trade are slow-growing and

difficult to propagate. Between 1997–2001, 99 percent of the total global trade in live corals

originated from “wild” sources and only 0.3 percent was captive-bred.

The economic viability of aquaculture must also be explored. In newly set-up operations, start-up

costs and operating costs are typically very high, with fairly low returns in comparison to wild-

caught products. However, these costs can be greatly reduced if established exporters set up

farming as a side industry. Governments and foreign aid may assist by providing initial funding.

To date, virtually all marine ornamentals fish are wild caught (breeding and rearing marine

species only accounts for 1–2 percent of the trade at present), and efforts to develop captive

cultivation have been limited.

Aquaculture can be an environmentally sound way to increase the supply of such organisms, by

helping reduce pressure on wild fish populations and producing juvenile and market-size fish of

a wide variety of species year-round. Furthermore, rearing aquarium fish in closed systems is

likely to lead to the production of hardier species, which fare better in captivity and survive

Blennies, gobies, and members of the Pomacentridae family are relatively easy to rear in

captivity as they attach or deposit their eggs on or in various substrates and, for species such as

the clown fish, can be conditioned to spawn voluntarily by manipulation of day length and water

temperature. Most other fish species such as angelfish and butterflyfish are known as broadcast

spawners, i.e., they spread their eggs freely in the water column and are therefore more difficult

to culture in captivity. They also usually require hormone treatment to induce spawning.

BIOL 181: Life in the Oceans – Lecture Notes

There is considerable interest in giant clam mariculture (Figure 25.9) for several reasons they

require no artificial feeding, rearing techniques are relatively simple, and the setting up of

facilities requires little capital investment and can involve local community members.

Furthermore, unlike many other forms of mariculture, it does not require broodstock to be

continuously captured from the wild, and hence the impact on wild stocks is minimal.

Giant Clam Mariculture, by JSLUCAS75, is available under a

Creative Commons Attribution-ShareAlike 3.0 Unported license.

Figure 25.9. Giant clam mariculture. Giant clams have been successfully grown in captivity.

Coral reefs are suffering from many threats, and it is important that the aquarium trade is

effectively managed in order to not further compound these problems. The United Nations

believes that if properly managed, the trade of marine ornamental species can help coastal

communities climb out of poverty.

The highly selective nature of this fishery increases the impact on populations of targeted

species. It may also, directly through the use of destructive fishing practices or indirectly through

the removal of key species (e.g., cleaner fish/shrimp), impact other species and ecological

processes in the habitats where fishing for the aquarium trade occurs.
BIOL 181: Life in the Oceans – Lecture Notes

25.9 Review Questions: Global Trade of Marine Ornamental Species

1. Is it just reef fish that are traded for aquaria?

2. True or false? The live coral trade for aquaria is significantly more valuable than sale to

the construction industry.

3. What is one of the main benefits of the aquarium trade for coral reef ecosystems?

4. How do they typically catch fish for the aquarium trade?

5. What are three problems associated with the aquarium trade?

6. What are two possible solutions to help alleviate problems associated with the aquarium


Profound inhibition of platelet aggregation with monoclonal antibody 7E3 Fab thrombolytic therapy : Results of the thrombolysis and angioplasty in mycardial infaraction (TAMI) 8 pilot study☆

Objectives. This study was undertaken to establish evidence for physiologic activity and to study the safety of murine-derived monoclonal antibody 7E3 Fab (m7E3 Fab) in receiving recombinant tissue-type plasminogen activator (rt-PA).

Background. Platelet aggregation is believed to be a significant factor in the failure of pharmacologic reperfusion. By binding to the glycoprotein IIb/IHa receptor, m7E3 Fab inhibits platelet aggregation and has been shown experimentally to decrease the time required for lysis and to prevent reocclusion. However, the safety of profound platelet inhibition after thrombolysis for acute myocardial infarction has not been tested in humans.

Methods. Sixty patients receiving rt-PA, aspirin and heparin for acute myocardial infarction received m7E3 Fab bolus injections in ascending doses at 3, 6 and 15 h after initiation of the thrombolytic infusion. Ten patients treated with rt-PA but not m7E3 Fab were studied as control subjects.

Results. Receptor site blockade and inhibition of platelet aggregation to 20 μmol/liter adenosine were maximal at a dose of 0.25 mg/kg body of m7E3 Fab. Fifteen (25%) m7E3 Fab-treated patients and five (50%) control patients had major bleeding eight of these events in seven m7E3 Fab-treated patients and one in a control patient occurred at the time of aortocoronary bypass surgery. Recurrent ischemia occured in eight (13%) m7E3 Fab-treated patients and two (20%) control subjects. Coronary angiography was performed in 43 patients the infarct-related coronary artery was patent in 5 of 9 (56%) control patients and 34 (92%) of 37 patients receiving m7E3 Fab.

Conclusions. Profound inhibition of platelet aggregation after thrombolysis was associated with bleeding rates comparable to those in control patients and a low rate of recurrent ischemia. The combination of m7E3 Fab and rt-PA, heparin and aspirin appears to be a promising and safe combination that should be evaluated in further studies of patients with acute myocardial infarction.

This study was supported by a grant from Centocor, Inc., Malvern, Pennsylvania. It was presented in part at the 64th Annual Scientific Sessions of the American Heart Association, Anaheim, California, November 1991.

#288 George Soros

Impact Investor The goal of impact investing is to create social or environmental impact, plus financial returns. The goal of impact investing is to create social or environmental impact, plus financial returns.

  • George Soros is a celebrated hedge fund tycoon who managed client money in New York from 1969 to 2011.
  • In 1992, Soros shorted the British pound and reportedly made a profit of $1 billion. He became known as the man who broke the Bank of England.
  • Soros shifted $18 billion from his family office to his Open Society Foundations as of 2018. His fund's assets are worth an estimated $8.4 billion.
  • Soros was born in Hungary at 17 he left the country and put himself through the London School of Economics working as a railway porter and waiter.
  • In recent public appearances, Soros has been a vocal critic of U.S. President Donald Trump and Chinese president Xi Jingping.

Did you know

Net worth over time

I occupy an exceptional position. My success in the financial markets has given me a greater degree of independence than most other people. This allows me to take a stand on controversial issues: in fact it obliges me to do so.

George Soros

Phils Sign OF Delmon Young To 1 Year Deal

Right-handed slugging outfielder was ALCS MVP last year, had 108 RBIs in 2010

By Todd Zolecki / | 01/22/2013 2:56 PM ET

PHILADELPHIA — The Phillies found a right-handed-hitting corner outfielder with power.

But this one comes with baggage.

The Phillies and free agent Delmon Young agreed Tuesday to a one-year contract worth $750,000 plus potential performance bonuses. The signing screams low-risk, high-reward for the Phillies. Young, who is 27, earned 2012 American League Championship Series MVP honors with the Detroit Tigers and hit .267 with 18 home runs and 74 RBIs in the regular season. He also had a career-high 112 RBIs in 2010 with the Minnesota Twins and has a career .824 OPS against left-handed pitchers.

But Young also has battled consistency issues on the field and has had several notable off-the-field problems, including an April arrest in New York on a second-degree aggravated harassment charge. Police said an intoxicated Young hurled anti-Semitic slurs at four men, and Major League Baseball suspended him for seven days as a result.

He was suspended twice in his Minor League career, for three games in 2005 for making contact with an umpire and for 50 games in 2006 for flinging a bat at an umpire.

However, the Phillies are comfortable with the risks, especially considering they made only a one-year commitment to him.

If Young plays well, the Phillies could have a productive corner outfielder at a remarkable bargain.

If he causes problems, well, the Phillies always can cut him loose without taking a significant payroll hit.

“Delmon is an experienced Major League bat who will add some depth to our relatively inexperienced outfield and another layer of competition for playing time there as well,” general manager Ruben Amaro Jr. said.

The Phillies had been looking for a corner outfielder since the beginning of the offseason. But how much outfield can Young play? And how effectively? He played just 31 games in the outfield last season, appearing in 113 games as Detroit’s designated hitter.

Young, listed at 6-foot-3 and 240 pounds, has had conditioning issues in the past, and last season he played with bone spurs in his ankle. He had surgery to remove the spurs in November and is expected to be ready for Spring Training.

Young’s arrival creates an interesting outfield situation for the Phillies. Ben Revere will be the team’s everyday center fielder. Young presumably will take one of the corner outfield spots, potentially as an everyday player. That could result in Domonic Brown and John Mayberry Jr. sharing time in the other corner spot with Laynce Nix a reserve and Darin Ruf possibly opening the season with Triple-A Lehigh Valley to get more seasoning.

Todd Zolecki is a reporter for This story was not subject to the approval of Major League Baseball or its clubs.

Delmon Damarcus Young

Position: Leftfielder
Bats: Right, Throws: Right
Height: 6′ 3″, Weight: 240 lb.

Born: September 14, 1985 in Birmingham, AL (Age 27)
High School: Adolfo Camarillo HS (Camarillo, CA)
by the Tampa Bay Devil Rays in the 1st round (1st pick) of the 2003 amateur draft.
Signed September 8, 2003. (All Transactions)
Debut: August 29, 2006 (Age 20)
Teams (by GP): Twins/DevilRays/Tigers 2006-2012
2013 Contract Status: Free Agent (details) [*]
Service Time (01/2013): 6.034, Free Agent: 2013 [*], Agents: Arn Tellem [*]
Relatives: Brother of Dmitri Young

1975 – 2012

Rk Year W L Ties W-L% Finish Playoffs R RA BatAge PitchAge Top Player Managers
30 1983 90 72 1 .556 1st of 6 Lost WS (4-1) 696 635 31.9 30.3 J.Denny (7.2) Corrales (43-42) & Owens (47-30)
31 1982 89 73 0 .549 2nd of 6 664 654 31.0 31.9 M.Schmidt (7.1) Pat Corrales (89-73)
32 1981 59 48 0 .551 3rd of 6 L LDS (3-2) 491 472 31.3 30.3 M.Schmidt (7.5) Dallas Green (59-48)
33 1980 91 71 0 .562 1st of 6 W WS (4-2) 728 639 30.6 28.6 S.Carlton (9.8) Dallas Green (91-71)
34 1979 84 78 1 .519 4th of 6 683 718 30.7 28.3 M.Schmidt (7.6) Ozark (65-67) & Green (19-11)
35 1978 90 72 0 .556 1st of 6 L NLCS (3-1) 708 586 29.9 30.3 M.Schmidt (6.0) Danny Ozark (90-72)
36 1977 101 61 0 .623 1st of 6 L NLCS (3-1) 847 668 29.4 29.7 M.Schmidt (8.7) Danny Ozark (101-61)
37 1976 101 61 0 .623 1st of 6 L NLCS (3-0) 770 557 28.8 29.8 M.Schmidt (7.9) Danny Ozark (101-61)
38 1975 86 76 0 .531 2nd of 6 735 694 27.7 27.3 M.Schmidt (7.4) Danny Ozark (86-76)


Exposure of male Fischer-344 (F-344) rats to methyl chloride (MeCI) results in testicular and epididymal toxicity and the induction of both pre- and postimplantation embryonic loss the preimplantation loss is caused by cytotoxic damage to sperm that leads to failure of fertilization (Toxicol Appl Pharmacol 1986 86: 124–130). The present study examined whether the cytotoxicity of MeCl to sperm is due to the testicular or epididymal toxicity of MeCl. Groups of 18 males were exposed to 3000 ppm MeCI 6 h/day for 5 days, with and without concurrent treatment with the anti-inflammatory agent 3-amino-1-[m-(trifluoromethyl)phenyl]-2-pyrazoline (BW755C 10 mg/kg, i.p. 1 h pre- and postexposure) BW755C was used to inhibit the epididymal toxicity of MeCI. Control groups were untreatd or injected as described above with BW755C. Six males from each group were killed weekly for 3 weeks. Toxic effects of MeCl on the testis were demonstrated by decreased relative organ weight (week 3), testicular histopathology (weeks 1–3) and decreased daily sperm production (weeks 1–3) these effects were not prevented by BW755C. In both the MeCI and the MeCI + BW755C treatment groups, tubules devoid of sperm were observed in regions 4 and 5 of the epididymis at week 2, and in regions 6A and 6B at week 3. Sperm were present in the vas deferens of both groups at week 3 in decreased numbers and had decreased motility and more frequent morphologic abnormalities compared to untreated controls. In conjunction with known epididymal transit times for F-344 rat sperm, these data indicate that the induction of preimplantation loss by MeCI at weeks 2 and 3 postexposure is likely to result from cytotoxic effects on sperm located in the testes at the time of exposure.

Are The Phils Headed For The Geriatric Ward?


They’re old. They’re broken down. They’re done.

The Philadelphia Phillies listened to the ridicule all month, as if their AARP cards are waiting in their mailbox, with retirement papers on the way.

“People keep talking about how old we are (30.8 average, second oldest in the major leagues) and how our window is about to close,” general manager Ruben Amaro says. “Maybe I’m delusional, but I really don’t think we’re old. We’re certainly not as old as other people think.

“I don’t believe our careers are over by any means.”

The Phillies might not be the same superpower that won five consecutive National League East titles, but after their 7-2 victory Wednesday against the Arizona Diamondbacks, they’re proving they’re not ready to surrender the throne.

The Phillies scored 20 runs in their last 19 innings against the Diamondbacks, and instead of answering questions about whether the end is near, they left town talking as if they’ve finally found themselves.

“The window closes every year, doesn’t it?” says Phillies shortstop Jimmy Rollins, 33. “We’ve had a long window, not as long as the (New York) Yankees or (Atlanta) Braves in their day, but the window closes every year because you have new personnel.

Philadelphia Phillies general manager Ruben Amaro Jr

“It’s different because of our personnel and the injuries we’ve had, but what makes it a lot different is that the reality is different than the perception. We may have to do things differently now, but we’re going to show the outside world that we still have plenty left in the tank.”

The Phillies, for years the Broad Street Bullies of the NL, suddenly look emaciated next to their former selves. They entered Wednesday having scored the third-fewest runs in baseball, and their 12 homers were three more than Los Angeles Dodgers center fielder Matt Kemp. They also had drawn the second-fewest walks, with two NL teams grounding into more double plays.

“We’re going to hear about our offense all year, but it’s going to be a different kind of offense that people are used to seeing,” leadoff hitter Juan Pierre said. “We’re not going to be sitting back hitting home runs. We’re going to scrape and scrap.”

They exemplified their sleeker selves Wednesday by amassing 13 hits (11 singles and two doubles).

The Phillies simply have no choice. They are without two-time home-run champ Ryan Howard (torn Achilles) for likely another month. All-Star second baseman Chase Utley (knee) still has no timetable for his return. They’re without 2008 Cy Young Award winner Cliff Lee, too.

“Every team has issues,” Amaro says. “Guys have to adjust.”

Rollins, normally the leadoff hitter, has two extra-base hits and three RBI as the No. 3 hitter. Right fielder Hunter Pence, normally the fifth-place hitter, has three hits in his last 24 at-bats as the cleanup hitter. And the Phillies are waiting for outfielder-first baseman John Mayberry Jr. (.200, two RBI) to simply hit.

“We have to keep plugging away and not get frustrated,” says starter Cole Hamels (3-1), who gave up four hits and two runs in eight innings and drove in two runs Wednesday. “We’re trying to play a different game now. Really, I think it’s just a matter of time.”

And when that time comes, the Phillies defiantly say, look out.

“We got knocked down,” Pence says. “We’ve got to keep getting back. And hopefully we get on a roll and start knocking other people down.”
– Copyright 2012 USA TODAY


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