M
Macroalgae: Diversity and
Conservation
Leonel Pereira
Department of Life Sciences, MARE – Marine
and Environmental Sciences Centre, Faculty of
Sciences and Technology, University of Coimbra,
Coimbra, Portugal
Definitions
Algae have a wide geographical distribution,
colonizing the most diverse sites, but always
associated with the presence of water. They
may be found floating in water, on wet rocks,
on walls, on other plants, or in association with
other organisms, as in the case of lichens, in
association with fungi. Likewise, most plants,
algae, have in their cells several pigments that
allow them to perform the photosynthesis and
can be fed by the absorption of organic sub-
stances present in the water.
The algae are abundant mainly in lakes, rivers,
and seas, occupying the euphotic (or photic)
region, that is, to the lower limit of luminous
penetration effective for algae photosynthesis
(up to about 100 m depth). Those living in water
may be planktonic (microalgae) or benthic
(macroalgae). The first ones live in suspension
and are of small size (microscopic); the latter are
© Springer Nature Switzerland AG 2022
W. Leal Filho et al. (eds.), Life Below Water, Encyclopedia of the UN Sustainable Development Goals,
https://doi.org/10.1007/978-3-319-98536-7
attached to the substrate (usually rocky) and are
larger (macroscopic), reaching 50 or more meters
in length. Any of various large brown seaweeds
(order Laminariales), which constitute the under-
water forests, are called “kelp.”
Introduction: Benthic Macroalgae
Macroalgae are aquatic photosynthetic organisms
belonging to the domain Eukarya and the king-
doms Plantae (green and red algae) and Chromista
(brown algae). The coloration of macroalgae rep-
resents the visible expression of multiple com-
bined pigments present in the cells; therefore,
phyla and classes are defined, in practice, by
their particular pigment composition (Pereira
2010a). Although classification systems are
diverse, it is generally agreed that (Fig. 1):
Green macroalgae are included in the phylum
Chlorophyta; their pigmentation is identical to
that of terrestrial plants (chlorophyll a, b, and
carotenoids).
Red macroalgae belong to the phylum
Rhodophyta and have as photosynthetic pigments
chlorophyll a, phycobilins (R-phycocyanin and
R-phycoerythrin), and carotenoids (β-carotene,
lutein, and zeaxanthin).
Brown macroalgae are included in the phylum
Heterokontophyta or Ochrophyta, in the class
Phaeophyceae; its pigments are chlorophyll a
528
Macroalgae: Diversity and Conservation, Fig. 1 The three main taxonomic groups of macroalgae: (a) Chlorophyta
(green algae); (b) Rhodophyta (red algae); (c) Phaeophyceae (brown algae)
and c and carotenoids where fucoxanthin predom-
inates, responsible for the brownish coloration.
Main Morphological Characteristics of
Benthic Macroalgae
The morphological diversity of algae is extraordi-
nary, in complexity, structure, and environmental
adaptations. Macroalgae exhibit a complex degree
of morphological organization, with laminar, cylin-
drical, tubular, or crustose thalli (Fig. 2). Some
algae are arranged as small thin disks, others as
fouling matter, or adhered to substrates. These thalli
are prostrated. However, most algae thalli are
upright (Fig. 2b–f), at least when immersed, during
the high tide period. The thallus of typical algae
(kelp, Fig. 3) is divided into a frond (the part that
stands erect) and a discrete fixation organ (as a
small disk or a tuft of thin elongated, colorless
elements) called rhizoids or holdfast. Only the
large (kelp) present a more robust fixation appara-
tus, composed of more or less curved elements, the
haptera (Pereira 2009; Braune and Guiry 2011;
Pereira and Correia 2015).
Filamentous thalli, the frond may be reduced
to a filament, where the cells are placed, linearly,
Macroalgae: Diversity and Conservation
one after the other. The filaments may be simple or
branched (in the form of a shrub). The ramifica-
tions can be irregular, dichotomous, alternating,
opposite, verticillate, or pectinate (Fig. 4).
Solid thalli have a compact structure, although
some species may have a soft consistency and
delicate texture. The solid thalli are subdivided
into four distinct types:
Cylindrical axes: may also be referred to by the
term “strands” provided they reach a considerable
diameter. These thalli are usually erect when
immersed, sometimes prostrating. They may be
branched according to the abovementioned way
(Fig. 4b).
Cylindrical or flat tubes: when the thalli are
hollow and their walls are composed of one or more
layers of cells. The axes of the tubes may constrict at
regular intervals (articulated stems) (Fig. 2d).
Blades: monostromatic (single layer of cells)
or polystromatic (two or more layers of cells). The
blades may be thin, more or less thick, or even
coriaceous and may be orbicular or elongated,
divided or not, and lobed or deeply divided (lacin-
iate or “belts”) (Fig. 2b).
The blades (or belts) can be traversed by “ribs”
(prominent, well-visible formations) or by
“veins” (queue of little prominent cells) (Fig. 2c).
Macroalgae: Diversity and Conservation 529

Macroalgae: Diversity and Conservation, Fig. 2 Morphological types: (a) crustose; (b) laminar; (c) erect; (d) hollow
tubular; (e) cylindrical; (f) vesicular

Vesicular: they are laminar stems, usually
polystromatic, with globose form (Fig. 2f).
Growth of thalli follow several modalities:
apical when the divisions are restricted to one or
more cells of the apical zone of the thallus; diffuse
when the divisions occur in different parts of the
stem; marginal (lamellar thalli) when only the
most peripheral cells divide regularly; and
meristodermal when a superficial layer of meri-
stematic cells found in some brown algae remains
active in the adult sporophyte and contributes to
intercalary growth.
530
Macroalgae: Diversity and Conservation,
Fig. 3 Thallus of a typical Kelp: A – Blade; B – Stipe;
C – Bulb; D – Papillae; E – Hapteron
Reproduction of Benthic Macroalgae
Reproduction of algae can be asexual and sexual.
Algae, both unicellular and multicellular, have
asexual and sexual reproduction. In single-cell
algae, asexual reproduction occurs through
binary division, with one individual giving rise
to two others. In the case of multicellular algae,
this type of reproduction can occur by fragmen-
tation of the thallus, as is common in filamentous
algae, or by zoosporia. In this process, an indi-
vidual form flagellated cells called zoospores
that loosen and, when fixed on some substrate,
originate new individuals. The sexual reproduc-
tion involves the fusion of male and female gam-
etes. The two types of reproduction often occur
in the same species, i.e., both sexual and asexual
generations coexist in the same individual. Only
the “blue algae” (Cyanobacteria) does not pre-
sent sexual reproduction; in this group there is
the transference of genetic material as in bacteria
(Braune and Guiry 2011; Pereira and Correia
2015).
Macroalgae: Diversity and Conservation
In vegetative and asexual reproduction, the
formation of new individuals can be done through
three distinct processes, which invest in homoge-
neity and promote the perpetuation of the genetic
heritage of the “progenitor”: (i) fragmentation of
thallus, where each small fragment grows into a
new thallus and a new individual; (ii) propagules,
i.e., through a cluster of cells, of small size, with
the ability to join a substrate and functionally
growth into to a new thallus; and (iii) spores,
which are cells formed inside sporocysts (special-
ized structures, originating from modified mother
cells).
The sexual reproduction represents the one in
which individuals invest substantially in speciali-
zation and in turn contribute to genetic diversity
from recombination (the parental characteristics
are somewhat diluted, assuming a compromise
between the genetic background of two different
individuals) (Fig. 5).
Underwater Forests of Benthic
Macroalgae (Kelp Forests)
Any diver to submerge on the rocky North Atlan-
tic coasts will easily find a unique landscape
consisting of algae that, up to 2 m high, take the
form of underwater forests. They are, as already
mentioned, the large brown seaweeds
(Phaeophyceae), whose density and large volume
allow them to host a splendid display of marine
animal life, of high ecological complexity, com-
parable only to deciduous forests in terrestrial
ecosystems. Its distribution starts from the begin-
ning of the infralittoral landing, with Saccorhiza
(Fig. 3), to continue in the following meters with
the Laminaria, until reaching the penetration of
the light, to more than 30 m of depth. But these are
not the only algae present in these forests, since
the protective conditions they offer, as well as the
creation of a gloomy environment, enable the
growth of a large variety of red, green, and
brown algae of small size, whose pigments allow
to take advantage of the luminous energy of less
intensity. The living beings that inhabit this eco-
system must choose, among the multiple habitats
available, the one that best suits their way of life.
Macroalgae: Diversity and Conservation 531

Macroalgae: Diversity
and Conservation,
Fig. 4 Branching types:
(a) irregular; (b)
dichotomous; (c)
alternating; (d) opposite;
(e) verticillate; (f) pectinate

From organisms that sessile with a benthonic
mode of life, or those that swim in the water
between the algae (nekton), like the fishes, not
forgetting those that are on the stipes and fronds
of the Laminaria (called epiphytes), or else refu-
gees in their rhizoids or haptera. All this gives us
an idea of the thousands of specimens we can find
in these forests. Of course, the “predators” of
these algae, the hedgehogs, cannot be lacking. It
is common to see clear openings in these forests,
completely devoid of Laminaria but full of hedge-
hogs, a situation which then takes advantage of
the fouling limestone algae (Lithophyllum
incrustans – Fig. 2a, Rhodophyta) to cover the
substrate, thus creating a habitat conducive to
colonization by anemones (phylum Cnidaria,
532
Macroalgae: Diversity and Conservation, Fig. 5 Two
types of fertilization can be differentiated in sexual repro-
duction in macroalgae: planogamy with both gametes are
mobile, and fertilization occurs between these mobile cells
(similar or not), after which they form a pair of swimmers
(planozygotes), during the period of their fusion in the
zygote; aplanogamy or oogamy with the male gametes
that goes against the female gametes and fuses with it. In
most cases, the female gametes are devoid of flagella and
are usually the one with the largest dimensions. In red algae
a particular type of oogamy occurs, as it happens between
different gametes, both not flagellated. In this case the
order Actiniaria). But if the hedgehogs control the
expansion of algae, then who controls the expan-
sion of their populations? The control is made by
fish and crustaceans, among which there are sev-
eral species that consider the hedgehogs a true
delicacy (Pereira and Correia 2015).
It is difficult to find in the rocky substratum an
uninhabited place, because the death of any sessile
organism (sponges, ascidians, mollusks, seaweed,
etc.) is used by another, of the same species or
different species, to seize this territory available.
Special mention should be made of the rhizoids and
haptera of Laminaria, since their multiple cavities
shelter a world of small organisms that find there
the ideal refuge. Sponges, sea lizards, mollusks,
echinoderms, anthozoans, and small fish are just a
few examples that give us an idea of the ecological
richness of these places. In this ecosystem the
Macroalgae: Diversity and Conservation
fertilization occurs in situ, that is, in the place where the
female gametes were formed. The species may be homo-
thallic (monoecious), i.e., both male and female sex organs
are present on the same thalli, or heterothallic (dioecious),
i.e., male and female sex organs develop on two different
thalli: A – spermatia (male gamete); B – trichogyne; C –
carpogonial branche; D – carpogonium (female gamete of
Florideophyceae, Rhodophyta); E – Fucus vesiculosus
thallus (Phaeophyceae); F – cross-section of a receptacle;
G – cross-section of a conceptacle; H – oosphere (male
gamete); I – anterozoid (male gamete)
whole substrate is good, even that of organic origin
that has already taken over, in the foreground, the
rocky substratum nude. Thus, the strains of Lami-
naria hyperborea present a roughness that allows
the adherence of algae, bryozoans, hydrozoans,
poriferous, among others which benefit from the
constant movement of water, forced to circulate
through this entanglement of life (and where the
water current loses force), to more easily capture
the suspended particles from which they feed.
Also, on the fronds it is easy to observe anthozoans,
such as anemones, or colonies of bryozoans that
even cover large surfaces (reminiscent of honey-
combs). About 13 species of kelp are known in
Atlantic European waters, one of them being non-
native, originating in Japan and introduced in
Europe (Undaria pinnatifida). Only recently the
existence of kelp has been confirmed in the
Macroalgae: Diversity and Conservation
archipelago of the Azores. The remaining species
are Saccorhiza polyschides, Laminaria
ochroleuca, L. hyperborea, Saccharina latissima,
Phyllariopsis brevipes, and P. purpurascens (Assis
et al. 2011).
Diversity and Conservation
The move toward a healthy ocean that is used
sustainably – the aim of Sustainable Development
Goal 14 – will assist in achieving several other
goals. Regarding macroalgae (seaweeds), goal 2
says the following: end hunger, achieve food
security and improved nutrition, and promote sus-
tainable agriculture. Ensuring proper management
of fish stocks and protecting fish and seaweed
quality will safeguard the vital role of food from
the sea in nourishing the world’s population
(United Nations 2017).
In 2005, world seaweed production totalled
14.7 million tonnes, of which the culture sector
contributed 13.5 million tonnes (freshwater cul-
ture, 53,157 tonnes; brackish water culture,
46,729 tonnes; and marine culture, 13.4 million
tonnes), while the harvest from the wild was
slightly over 1.2 million in the same year. Ten
years later, by 2015, the total production had
doubled to 30.4 million tonnes, with the culture
and capture sectors responsible for 29.4 million
tonnes and 1.1 million tonnes, respectively. The
leading producers by country (in order of decreas-
ing rank) were Chile, China, and Norway for wild
species (mainly brown and red algae) and China,
Indonesia, Republic of Korea, and the Philippines
for cultured species (mainly Eucheuma/
Kappaphycus, Saccharina japonica, Gracilaria,
and Undaria pinnatifida) (FAO 2018).
Climate Change and Ocean Acidification
Seaweed and seagrass communities in the North-
east Atlantic have been profoundly impacted by
humans, and the rate of change is accelerating
rapidly due to runaway CO2 emissions and
mounting pressures on coastlines associated with
human population growth and increased con-
sumption of finite resources. Here, we predict
533
how rapid warming and acidification are likely
to affect benthic flora and coastal ecosystems of
the Northeast Atlantic in this century, based on
global evidence from the literature as interpreted
by the collective knowledge of the authors. Brodie
et al. (2014) predict that warming will kill off kelp
forests in the south and that ocean acidification
will remove maerl habitat in the North Atlantic
(Fig. 6). Maerl (or rhodolith) is mainly composed
of three species of coralline algae: Lithothamnion
corallioides, L. glaciale, and Phymatolithon
calcareum (Irvine and Chamberlain 1994; Guiry
and Guiry 2020). Maerl is dredged from the sea-
floor and crushed to form a powder. It is still
harvested around the coasts of Brittany in France
and Bantry Bay, Ireland, and is a popular fertilizer
for organic gardening (Blunden et al. 1981).
Seagrasses will proliferate, and associated epi-
phytes switch from calcified algae to diatoms and
filamentous species. Invasive species will thrive
in niches liberated by loss of native species and
spread through exponential development of artifi-
cial marine structures (Brodie et al. 2014). Com-
bined impacts of seawater warming, ocean
acidification, and increased stormy conditions
M
may replace structurally diverse seaweed cano-
pies and associated calcified and noncalcified
flora with simple habitats dominated by non-
calcified, turf-forming seaweeds (Cambridge
et al. 2007; Brodie et al. 2014).
On the other hand, where waters remain cool
enough, assemblages of fleshy macroalgae are
expected to benefit from high CO2 conditions as
increased inorganic carbon availability may
enhance the growth and reproduction of fleshy
macroalgae (Harley et al. 2012; Koch et al.
2013; Kroeker et al. 2013). Figure 6 shows several
examples of how such changes are predicted to
affect the Northeast Atlantic where the flora is
dominated by kelps (Laminariales) in the subtidal
and fucoids (Fucales) in the intertidal (Brodie
et al. 2014).
Greenhouse Effect, Air Purification, and
Recycling of Organic Matter
Through photosynthesis, algae, like any other
plant, on one hand, fix and subtract atmospheric
CO2, contributing to the reduction of its concen-
tration (related and contributory causal agent to
534
Macroalgae: Diversity and Conservation,
Fig. 6 Estimation of the effects of increased CO2 and
consequent acidification of the oceans on the North Atlan-
tic marine flora: (a) In Artic region the warming will be
detrimental to cold-adapted species, and acidification will
corrode maerl (M). Pacific species, like Alaria marginata
(Am), will invade as polar ice melts, competing with native
Macroalgae: Diversity and Conservation
species such as Laminaria hyperborea (Lh) and Alaria
esculenta (Ae). Fleshy invasive, for example, Sargassum
muticum (Sm), will move north competing with fucoids,
for example, Fucus distichus (Fd), in the intertidal zone.
Acidification will corrode epiphytic calcified algae, for
example, Titanoderma pustulatum (Tp), and increased
CO2 levels will stimulate growth of diatoms (D) (drawings
Macroalgae: Diversity and Conservation
the “greenhouse effect,” which causes a progres-
sive increase in temperature of the planet), and, on
the other hand, release O2, gas consumed by all
living beings in the process of cellular respiration
which is essential to produce chemical energy.
Thus, algae are fundamental in the process of
purifying the air that we breathe while helping to
recycle the raw materials (Pereira and Correia
2015).
Marine Algae Aquaculture: Promoting Both
Carbon Sequestration and Sustainability
Current atmospheric CO2 levels (400 ppm) are
well above those considered safe for the preserva-
tion of the planet and supporting life on Earth.
This was established by researchers as 350 ppm
(Rockstrom et al. 2009); see www.350.org for
further information. Proactive policies that com-
bine actions to remove CO2 from the atmosphere
with actions to reduce emissions could decrease
CO2 concentrations faster than possible via natu-
ral processes (Lemoine et al. 2012). Among these
carbon-negative actions is the bioenergy with car-
bon capture (BEBCCS) which combines biologi-
cal carbon capture and sequestration with biomass
production (Mathews 2008). Because of their
very high productivity, marine algae offer great
potential for carbon-negative bioremediation
approaches (Arenas and Vaz-Pinto 2015).
Macroalgae Carbon Sequestration
For centuries, humans have harvested seaweed for
food and chemical constituents. Nowadays, as
food and energy demand continues to rise, marine
macroalgae are receiving increasing attention as
an attractive renewable source of food, fuels, and
other chemicals. Today, the world seaweed
Macroalgae: Diversity and Conservation, Fig. 6 (con-
tinued) made under the microscope) and marine angio-
sperms such Zostera marina (Zm). (b) In Boreal region,
Laminaria Hyperborea (Lh) forests will be increasingly
dominated by Laminaria ochroleuca (Lo), with the loss
of Alaria esculenta (Ae) and fucoids, for example, Fucus
vesiculosus (Fv) and the continued spread of invasive
Undaria pinnatifida (Up), Sargassum muticum (Sm), and
Grateloupia turuturu (Gt). As in the Artic, maerl beds will
be corroded, marine angiosperms will thrive, but epiphytic
535
industry is worth more than US$ billion per year
(which corresponds to approximately 12 million
tons per year), of which about 85% are food
products for human consumption (Arenas and
Vaz-Pinto 2015). Extracts derived from macro-
algae (alginates, carrageenan, and agar) represent
almost 40% of the world’s hydrocolloid market in
terms of food; the rest comes from certain ani-
mals, microbes, and land plants (FAO 2018).
Macroalgae contain an average of 30% carbon,
so this figure represents 4.71
106 tons of carbon
harvested from macroalgae annually (Arenas and
Vaz-Pinto 2015). About 221 species of macro-
algae have commercial value, and about 10 spe-
cies are cultivated in an intensive way, as is the
example of the brown algae (Undaria pinnatifida
(Harvey) Suringar, Saccharina japonica
(Areschoug) C.E. Lane, C. Mayes, Druehl & G.
W. Saunders, and Sargassum fusiforme (Harvey)
Setchell), red algae (Pyropia/Porphyra spp.,
Kappaphycus alvarezii (Doty) Doty ex P.C.
Silva, Eucheuma spp., and Gracilaria spp.), and
green algae (Monostroma nitidum Wittrock, Ulva
clathrata (Roth) C. Agardh, and Caulerpa spp.).
The cultivation of Japanese kelp (Saccharina
M
japonica) corresponds to about 33% of the total
cultivated macroalgae (FAO 2018). Macroalgal
farms may be integrated as offshore farms, near-
shore farms, and land-based ponds. There is evi-
dence that current culture production is enough to
meet the needs of the food and commodity mar-
kets (Roesijadi et al. 2010). However, recent
advances in the technology to convert carbohy-
drates from seaweeds into biofuels have awak-
ened the interest in improving seaweed
cultivation methods. Worldwide energy demand
is increasing, and there is a growing dilemma
calcified algae will be reduced or replaced with diatoms
(D) and filamentous algae. (c) In Lusitanian region, kelps
(Laminaria ochroleuca – Lo, and Saccorhiza polyschides –
Sp) will be replaced by smaller, fleshy algae and invasive
species, for example, Caulerpa taxifolia (Ct) will prolifer-
ate. Fucoids will be replaced by invasive such as
Asparagopsis armata (Aa). Marine angiosperms will
thrive, and it is expected that maerl and epiphytic calcified
algae will be retained (adapted from Brodie et al. 2014)
536
about finding a feedstock capable of keeping up
with demand. Currently, one fifth of the global
CO2 emissions are from the transport sector
(Rawat et al. 2013). The continued use of fossil
fuels is not sustainable, and biofuel production,
i.e., derived from biological carbon sequestration,
has emerged as an effective alternative (Demirbas
2007; Singh et al. 2011). The use of renewable
biomass as an energy resource has good benefits
to the environment (lower toxicity and lower
emission profiles). Biodiesel and bioethanol syn-
theses from terrestrial food crops are attractive
and widely used because of well-established farm-
ing practices and simple and cheap extraction
processes (Wei et al. 2013). However, conven-
tional biofuels from classic food crops require
high-quality agricultural land, and there are large
concerns on future competition with food agricul-
ture for arable land use (Goh and Lee 2010). One
plausible alternative to food crops is the use of
nonfood lignocellulosic biomass such as agricul-
tural residues or wood waste. However, current
technologies have yet to solve the problems
related with sugar extraction from these products,
namely, from algal biomass (Wei et al. 2013;
Arenas and Vaz-Pinto 2015; Pereira 2017).
Novel Applications of Seaweeds
Since ancient times, seaweeds are a direct food
source for humans, especially in East Asia
(Japan), the Indo-Pacific (China, Indochina), and
Pacific (Hawaii). In China and Japan, more than
70 species of marine macroalgae are consumed,
and in both countries, there are marine farms with
hundreds of square km of area, where specific
species (Porphyra/Pyropia for “Nori,”
Saccharina/Laminaria for “Kombu,” Undaria
for “Wakame”) are cultivated for human con-
sumption (Braune and Guiry 2011; Pereira 2016).
One of the most suitable strategies is the inte-
grated multi-trophic aquaculture system (IMTA).
This situation resolves the nutrient issue in more
than one way. As mentioned elsewhere, fish aqua-
culture causes the problem of high nutrient pro-
duction due to fish or shellfish feeding and
excrement. If these waters get to a seaweed
Macroalgae: Diversity and Conservation
production tank before reaching the environment
again, the nutrient uptake by seaweed will solve
the high nutrient pollution and stimulate the bio-
mass production of algae (Pinto and Abreu 2011;
Abreu et al. 2011).
Now, for the industrial part, the best way to
profit from seaweed is to use all its content or at
least attempt to do so. Normally, a company
focuses on a single compound of interest; how-
ever, it would be desirable to consider the
remaining contents of the seaweed. We know,
for instance, that some seaweeds are used as
food, but they also have potential health and
body-care properties, and that they are rich in
sugars, vitamins, minerals, and proteins – in
other words, they are rich sources of useful
products.
Thus, it is in a company’s best interests to
consider not only the compound selected for its
high market value but all the other compounds as
well. Algae are a very good resource as fertilizers
for agriculture and vegetal solution for animal
feeding such as pet and cattle. They are source
of many compounds for the cosmetic, food, and
pharmaceutical industries. Their notable sugar
content is a profitable way to produce bioethanol
through the seaweed biomass residues produced
from another industry.
Sometimes the aggressive methods used in the
industry do not allow the efficient use of the total
content of the biomass, but with a carefully
designed method, we can improve the efficiency
and obtain more than just one compound/product
(Carvalho and Pereira 2015).
Historically, the algae have been used by
coastal communities to prepare home medicines
to treat various health problems. Such applica-
tions are the product of the traditional knowledge
of many generations, and, in most cases, their
mechanisms are unknown. So, we know, for
example, that the good results obtained with the
use of kelp (Laminaria spp.) in the treatment of
goiter are due to the presence of iodine in brown
algae (Pereira 2010b). Green algae (Chlorophyta)
have been used as anthelmintic and astringent and
to treat gout. Brown algae are used in the treat-
ment of rheumatic processes, arteriosclerosis,
menstrual disorders, hypertension, gastric ulcers,
Macroalgae: Diversity and Conservation
goiter, skin diseases, syphilis, and as an anticoag-
ulant. Red algae are used as anticoagulants,
anthelmintic, and in treating gastritis and diarrhea
(Ebadi 2007; Pereira 2011, 2018).
An important issue pertaining to the sustain-
able future for the world algae consumption is the
potential of algae cultivation equivalent to terres-
trial agriculture. In Portugal, for example, sustain-
able production of seaweed and seaweed-based
products (see www.algaplus.pt) involves the inte-
grated multi-trophic aquaculture (IMTA) concept
– farming seaweeds in proximity to several spe-
cies at different trophic levels, allowing the reduc-
tion of aquaculture wastes (Troell et al. 2009).
IMTA can be practiced in a controlled environ-
ment on land, with organic certification for qual-
ity, traceability, stability of supply, and a small
carbon footprint (Buschmann et al. 2017).
An expanding body of evidence suggests that
offshore-cultivated seaweeds, which do not com-
pete with food crops for arable land or drinkable
water, could provide an alternative source of bio-
mass for the sustainable production of food,
chemicals, and fuels (Radulovich et al. 2015;
Neori 2016). However, increasing the global pro-
duction of seaweeds requires an understanding of
the critical points that currently limit their produc-
tion. Some studies developed models for
assessing the net primary productivity
(Buschmann et al. 2017). The scale of terrestrial
agriculture beats seaweed aquaculture by two
orders of magnitude (Neori et al. 2016). Adjusting
this immense production inequity is an intimidat-
ing challenge (Neori 2016; Neori et al. 2016).
Nevertheless, several authors claim that seaweed
aquaculture can produce several billion tones
year1 of macroalgae, which would provide a
maintainable supply of affordable and healthy
food for the centuries to come (Lenstra et al.
2011; Radulovich et al. 2015; Bjerregaard et al.
2016; Kim et al. 2017). Such large-scale efforts in
mariculture might be located on the surface of the
oceans and in coastal deserts, using seawater with
natural nutrients or aquaculture waste. Raising
industrial output by two orders of scale would
require a system of aquaculture that can be oper-
ated at an unprecedented scale while meeting the
environmental, social, and economic goals of
537
sustainable development (Bjerregaard et al.
2016), just as land plants led agriculture
(Buschmann et al. 2017).
Many products we consume or use in daily life
contain seaweed extracts in some form such as
ham, ice cream, bottled chocolate drinks, and
toothpaste or deodorant, although most people
probably don’t even realize it. There is currently
a growing interest in algae due to the recognition
of numerous new bioactive compounds. Antioxi-
dants, antimicrobials, anti-inflammatory, anti-
aging, and antitumor are just some of its
amazing properties for use as pharmaceuticals,
cosmeceuticals, nutraceuticals, or even in agricul-
ture or food (Leandro et al. 2020). There is an
increasing awareness of the sustainable use of
natural resources rather than synthetic and pro-
cessed products with possible harmful consumer
side effects. The growing interest in these poten-
tialities has led to the promotion of macroalgae
production, as well as to new topics of research.
This augurs well for future development of this
industry on a sustainable basis.
M
Cross-References
▶ Biological Invasions as a Threat to Global
Sustainability
▶ Marine Ecosystems
▶ Ocean Sustainability
▶ Resilient Oceans: Policies and Practices to Pro-
tect Marine Ecosystems
▶ Trends and Patterns of the Seaweed Industry
and Its Links with SDGs
References
Abreu MH, Pereira R, Mata L, Nobre A, Pinto IS (2011)
Aquacultura Multi-Trófica Integrada em Portugal. In:
Ferreiro UV, Filgueira MI, Otero RF, Leal JM (eds)
Macroalgas en la Acuicultura Multitrófica Integrada
Peninsular. Valorización de su Biomasa. Centro
Tecnológico del Mar – Fundación CETMAR, Spain,
Vigo, pp 54–77. ISBN: 978-84-615-4974-0
Arenas F, Vaz-Pinto F (2015) Chapter 5 – marine algae as
carbon sinks and allies to combat global warning. In:
Pereira L, Neto JM (eds) Marine algae – biodiversity,
taxonomy, environmental assessment and
538
biotechnology. CRC Press, Taylor & Francis Group,
Boca Raton. ISBN 978-1-46658-167-8
Assis J, Tavares JT, Serrão EA, Alberto F, Ferreira C,
Tavares D, Paulos L, Tempera F (2011) Florestas
Marinhas – As Espécies de Algas Castanhas Gigantes
de Portugal. Mundo Gobius Comunicação e Ciência,
Lda., 32 p. ISBN 978-989-97260-0-0
Bjerregaard R, Valderrama D, Radulovich R, Diana J,
Capron M, Mckinnie CA, Cedric M, Hopkins K, Yarish
C, Goudey C, Forster J (2016) Seaweed aquaculture for
food security, income generation and environmental
health in tropical developing countries. World Bank
Group, Washington, DC. Available online at: http://
docum ents.worldbank.org/curated/pt/947831
469090666344/Seaweed-aquaculture-for-food-security-
income-generation-and-environmental-health-in-Tropi
cal-Developing-Countries
Blunden G, Farnham WF, Jephson N, Barwell CJ, Fenn
RH, Plunkett BA (1981) The composition of
Maerl beds of economic interest in northern Brittany,
Cornwall, and Ireland. Proc Int Seaweed Symp
10:651–656
Braune W, Guiry M (2011) Seaweeds – a colour guide to
common benthic green, brown and red algae of the
world’s oceans. A.R.G. Gantner Verlag K.G, Ruggell.
ISBN 978-3-906166-90-2
Brodie J, Williamson CJ, Smale DA, Kamenos NA,
Mieszkowska N, Santos R, Cunliffe M, Steinke M,
Yesson C, Anderson KM, Asnaghi V, Brownlee C,
Burdett HL, Burrows MT, Collins S, Donohue PJ,
Harvey B, Foggo A, Noisette F, Nunes J, Ragazzola
F, Raven JA, Schmidt DN, Suggett D, Teichberg M,
Hall-Spencer JM (2014) The future of the Northeast
Atlantic benthic flora in a high CO2 world. Ecol Evol 4
(13):2787–2798. https://doi.org/10.1002/ece3.1105
Buschmann AH, Camus C, Infante J, Neori A, Israel A,
Hernández-González MC, Pereda SV, Gomez-Pinchetti
JL, Golberg A, Tadmor-Shalev N, Critchley AT (2017)
Seaweed production: overview of the global state of
exploitation, farming and emerging research activity.
Eur J Phycol 52(4):391–406. https://doi.org/10.1080/
09670262.2017.1365175
Cambridge ML, How JR, Lavery PS, Vanderklift MA
(2007) Retrospective analysis of epiphyte assemblages
in relation to seagrass loss in a eutrophic coastal embay-
ment. Mar Ecol Prog Ser 346:97–107. https://doi.org/
10.3354/meps06993
Carvalho LG, Pereira L (2015) Review of marine algae
as source of bioactive metabolites. In: Pereira L,
Neto JM (eds) Marine algae – biodiversity, taxon-
omy, environmental assessment and biotechnology.
Science Publishers, an Imprint of CRC Press, Taylor
& Francis Group, Boca Raton. ISBN 978-1-46658-
167-8
Demirbas A (2007) Alternatives to petroleum diesel fuel.
Energy Sources 2(4):343–351. https://doi.org/10.1080/
15567240600629518
Ebadi MS (2007) Pharmacodynamic basis of herbal med-
icine. CRC Press, Taylor and Francis Group, Boca
Raton. ISBN 978-0849370502
Macroalgae: Diversity and Conservation
FAO (2018) The global status of seaweed production.
Globefish research programme, vol 124. FAO, Rome,
120 p. ISBN 978-92-5-130870-7
Goh CS, Lee KT (2010) A visionary and conceptual
macroalgae-based third-generation bioethanol (TGB)
biorefinery in Sabah, Malaysia as an underlay for
renewable and sustainable development. Renew Sus-
tain Energy Rev 14(2):842–848. https://doi.org/10.
1016/j.rser.2009.10.001
Guiry MD, Guiry GM (2020) AlgaeBase. World-wide
electronic publication, National University of Ireland,
Galway. Available online at: http://www.algaebase.org
Harley CDG, Anderson KM, Demes KW, Jorve JP, Kordas
RL, Coyle TA, Graham MH (2012) Effects of climate
change on global seaweed communities. J Phycol
48:1064–1078. https://doi.org/10.1111/j.1529-8817.2012.
01224.x
Irvine LM, Chamberlain YM (1994) Seaweeds of the Brit-
ish Isles. volume 1, Part 2B. The Natural History
Museum, London. ISBN 0-11-310016-7
Kim JK, Yarish C, Hwang EK, Park M, Kim Y (2017)
Seaweed aquaculture: cultivation technologies, chal-
lenges and its ecosystem services. Algae 32:1–13.
https://doi.org/10.4490/algae.2017.32.3.3
Koch M, Bowes G, Ross C, Zhang X-H (2013) Climate
change and ocean acidification effects on seagrasses
and marine macroalgae. Global Change Biology 19
(1):103–132. https://doi.org/10.1111/j.1365-2486.
2012.02791.x
Kroeker KJ, Kordas RL, Crim R, Hendriks IE, Ramajo L,
Singh GS, Duarte CM, Gattuso J-P (2013) Impacts of
ocean acidification on marine organisms: quantifying
sensitivities and interaction with warming. Global
Change Biology 19(6):1884–1896. https://doi.org/10.
1111/gcb.12179
Leandro A, Pereira L, Gonçalves AMM (2020) Diverse
applications of marine macroalgae. Mar Drugs 18
(1):17. https://doi.org/10.3390/md18010017
Lemoine DM, Fuss S, Szolgayova J, Obersteiner M,
Kammen DM (2012) The influence of negative
emission technologies and technology policies on the
optimal climate mitigation portfolio. Clim Chang
113(2):141–162. https://doi.org/10.1007/s10584-011-
0269-4
Lenstra J, Hal VJ, Reith H (2011) Economic aspects of open
ocean seaweed cultivation. ECN – Energy Research
Centre of the Netherlands. Montpellier, France. Avail-
able online at: http://marineagronomy.org/sites/default/
files/Economics%20aspects%20of%20open%20ocean
%20seaweed%20cultivation.pdf
Mathews JA (2008) Carbon-negative biofuels. Energy Pol-
icy 36(3):940–945. https://doi.org/10.1016/j.enpol.
2007.11.029
Neori A (2016) Can sustainable mariculture match agricul-
ture’s output? Global Aquaculture Alliance. Available on-
line at: https://www.aquaculturealliance.org/advocate/
can-sustainable-mariculture-match-agricultures-output/
Neori A, Shalev TN, Agami M (2016) Seaweed aquacul-
ture as a match for agriculture. In: 22nd international
seaweed symposium, Copenhagen
Management and Monitoring of Eutrophication
Pereira L (2009) Guia Ilustrado das Macroalgas –
Conhecer e Reconhecer Algumas Espécies da Flora
Portuguesa. Imprensa da Universidade de Coimbra,
Coimbra. ISBN 978-989-26-0002-4
Pereira L (2010a) Littoral of Viana do Castelo – ALGAE
(Bilingual). Câmara Municipal de Viana do Castelo.
ISBN: 978-972-588-217-7
Pereira L (2010b) Littoral of Viana do Castelo – ALGAE:
agriculture, gastronomy and food industry (Bilingual).
Câmara Municipal de Viana do Castelo. ISBN 978-
972-588-218-4
Pereira L (2011) A review of the nutrient composition of
selected edible seaweeds. In: Pomin VH (ed) Seaweed:
ecology, nutrient composition and medicinal uses.
Nova Science Publishers Inc., New York. ISBN: 978-
1-61470-920-6
Pereira L (2016) Edible seaweeds of the world. Science
Publishers, an Imprint of CRC Press, Taylor & Francis
Group, Boca Raton. ISBN 978-1498730471
Pereira L (ed) (2017) Algal biofuels. Science Publishers, an
Imprint of CRC Press, Taylor & Francis Group, Boca
Raton. ISBN 978-1498752312
Pereira L (2018) Therapeutic and nutritional uses of algae.
Science Publishers’ (SP), An Imprint of CRC Press,
Taylor & Francis Group, Boca Raton. ISBN 978-
1498755382
Pereira L, Correia F (2015) Algas Marinhas da Costa
Portuguesa – Ecologia, Biodiversidade e Utilizações.
Nota de Rodapé Edições. ISBN 978-989-20-5754-5
Pinto IS, Abreu H (2011) Aquacultura Multitrófica
Integrada – O que é? In: Ferreiro UV, Filgueira MI,
Otero RF, Leal JM (eds) Macroalgas en la
Acuicultura Multitrófica Integrada Peninsular.
Valorización de su Biomasa. Centro Tecnológico
del Mar – Fundación CETMAR, Spain. Vigo,
ISBN: 978-84-615-4974-0
Radulovich R, Neori A, Valderrama D, Reddy CRK, Cro-
nin H, Forster J (2015) Farming of seaweeds. In: Tiwari
B, Troy D (eds) Seaweed sustainability – food and non-
food applications. Academic Press, London. https://
doi.org/10.1016/B978-0-12-418697-2.00003-9
Rawat I, Kumar RR, Mutanda T, Bux F (2013) Biodiesel
from microalgae: A critical evaluation from laboratory
to large scale production. Applied Energy 103:444–
467. https://doi.org/10.1016/j.apenergy.2012.10.004
Rockstrom J, Steffen W, Noone K, Persson V, Chapin FS,
Lambin EF, Lenton TM, Scheffer M, Folke C,
Schellnhuber HJ, Nykvist B, de Wit CA, Hughes T,
van der Leeuw S, Rodhe H, Sörlin S, Snyder PK,
Costanza RU, Svedin M, Falkenmark L, Karlberg
RW, Corell VJ, Fabry J, Hansen B, Walker D, Liverman
D, Richardson K, Crutzen P, Foley JA (2009) A safe
operating space for humanity. Nature 461(7263):472–
475. Available online at: https://www.nature.com/
articles/461472a.pdf
Roesijadi G, Jones SB, Snowden-Swan LJ, Zhu Y (2010)
Macroalgae as a biomass feedstock: a preliminary anal-
ysis. In: U.S.D.O. Energy: Pacific Northwest National
Laboratory (ed), 19944. Available online at: http://
sailing-sea-farm.com/onewebmedia/PNNL-19944.pdf
539
Singh A, Nigam PS, Murphy JD (2011) Renewable fuels
from algae: an answer to debatable land based fuels.
Bioresour Technol 102(1):10–16. https://doi.org/10.
1016/j.biortech.2010.06.032
Troell M, Joyce A, Chopin T, Neori A, Buschmann AH, Fang
JG (2009) Ecological engineering in aquaculture – poten-
tial for integrated multi-trophic aquaculture (IMTA) in
marine offshore systems. Aquaculture 297:1–9. https://
doi.org/10.1016/j.aquaculture.2009.09.010
United Nations (2017) The ocean and the sustainable
development goals under the 2030 agenda for sustain-
able development. A technical abstract of the first
global integrated marine assessment. UN, New York,
48 p. eISBN 978-92-1-361371-9
Wei N, Quarterman J, Jin YS (2013) Marine macroalgae:
an untapped resource for producing fuels and
chemicals. Trends Biotechnol 31(2):70–77. https://
doi.org/10.1016/j.tibtech.2012.10.009
Management and Monitoring
of Eutrophication: Trophic
State Indexes on the Río de la
Plata Northern Coast
Ernesto Brugnoli1,2, Rafael Arocena3,
Lucía Cabrera-Lamanna4 and Pablo Muniz1
1 M
Oceanografía y Ecología Marina, Facultad de
Ciencias, Universidad de la República,
Montevideo, Uruguay
2
Estación Científica COIBA-AIP, Clayton,
Panamá
3
Sección Limnología, Facultad de Ciencias,
Universidad de la República, Montevideo,
Uruguay
4
Departamento de Ecología y Gestión Ambiental,
Centro Universitario Regional Este, Universidad
de la República, Maldonado, Uruguay
Synonyms
Coastal environmental quality; Ecosystem health;
Estuary; Organic enrichment
Definitions
Marine pollution is an introduction by humans,
directly or indirectly, of substances or energy into
marine environments (including estuaries) resulting