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Physiological Characteristics of Porphyra
RUI PEREIRA AND CHARLES YARISH
Some species of Porphyra occur in the intertidal (e.g., P. dioica, P. umbilicalis,
P. linearis) while others are found only in the low-intertidal (e.g., P. leucosticta, P. purpurea) or the subtidal regions (e.g. P. amplissima). Intertidal species are
naturally expected to survive salinity variations, but in reality not many studies
have looked into the effects of salinity in Porphyra. Stekoll et al. (1999) investigated
the effects of salinity on the growth of the conchocelis phase of P. abbottae, P. torta
and P. pseudolinearis. Those authors found that salinity between 20% and 40‰
had little effect on growth, but there was virtually no growth at salinity 10‰ and
below. Conitz et al. (2001), on the other hand, did not find significant effects of
salinity on growth of juvenile gametophytes of P. torta, from 30‰ to 7.5‰.
The effects of photon flux density (PFD) and photoperiod have been studied
for many species, especially due to their influence in the life cycle of virtually all
species. The gametophytes and sporophytes of Porphyra dioica, for instance, grow
in short-day, neutral-day and long-day photoperiods. However, formation of
conchosporangia and release of conchospores in the laboratory occurred only in
short days (Pereira et al., 2006).
In terms of nutrient removal, the works by Chopin et al. (1999), Carmona
et al. (2006) and Pereira et al. (2008), among others, showed that some species of
Porphyra perform well in that function. Chopin et al. (1999) showed that tissue
nitrogen (N) and phosphorus (P) of Porphyra increased in specimens grown near
salmon cages. Of the several species studied, these authors concluded that
Porphyra yezoensis and Porphyra purpurea respond to high nutrient loading in
coastal waters resulting from anthropogenic activities (salmon aquaculture and
intense scallop dragging).
Studies in the laboratory have also shown that Porphyra responds to high
concentrations of ammonium (NH4+), a common form of N in water enriched by
anthropogenic activities. Wu et al. (1984) studied the utilisation of NH4+ by
Porphyra yezoensis and obtained higher growth rates (11.6% day−1) and tissue nitrogen content (4.72% dry weight [dw]) with NH4+ concentration ranging from 5 to 10
ppm (approx. 350–700 µM). For the same species, Amano and Noda (1987)
reported that the optimal fertilising effect was obtained using 20 ppm NH4+ during
48 h. More recently, Carmona et al. (2006) assessed the bioremediation potential of
several native northeast American species of Porphyra and compared those with
well-known Asian species. In that study, growth and tissue N reached maximal
levels at inorganic N concentrations of 150–300 µM. Maximum growth rates
ranged from 10 to 25% day−1. Pereira et al. (2008) studying a North Atlantic species,
Porphyra dioica, also reported an interesting capacity to uptake equally well NO3−
or NH4+, in concentrations ranging from 25 to 300 µM, when only one of these N
forms is present. If both forms were present, the preference was for NH4+ even if
available at a much lower concentration than NO3−.
Still in terms of nutrient physiology, considerably fewer studies have focused on
aspects related to phosphorus (P) and carbon (C). Carmona et al. (2006) showed that
P biofiltering efficiency was higher when NH4+ was supplied in P. amplissima,
P. purpurea, P. umbilicalis and P. yezoensis, but not in P. katadai and P. haitanensis.
THE ROLE OF PORPHYRA IN SUSTAINABLE CULTURE SYSTEMS
These authors also determined that P uptake was not saturated up to 30 µM P in
P. umbilicalis and P. haitanensis, while uptake rate by P. purpurea was saturated at
15 mM P, regardless of the N source, and at even lower P concentrations in the
other species. Pereira et al. (2008) showed that, for P. dioica, P enrichment up to
400 µM P did not translate into higher P uptake. In fact, despite the increasing P
concentration, the total amount of P removed from the medium was the same.
Both studies confirm what was first pointed out by Hafting (1999) for P. yezoensis,
a lack of capacity to store P. Interestingly, Chopin et al. (2004) described the presence of polyphosphate granules in the cells of P. purpurea. Polyphosphate granules are a form of P storage in yeast, microalgae and some macroalgae Sommer
and Both, 1938; Lundberg et al., 1989; Chopin et al., 1997). Chopin et al. (2004),
however, also point out that these granules do not seem to have the same storage
function in P. purpurea.
In terms of C metabolism, there are seaweed species capable of using only
CO2 and others that can use CO2 or HCO3−. Bicarbonate utilisation has been suggested for P. leucosticta (Mercado et al., 1997), P. umbilicalis (Maberly, 1990) and,
although with a limited capacity, for P. linearis (Israel et al., 1999). For P. leucosticta and P. linearis, those authors detected the activity of intra- and extracellular
carbonic anhydrase (CA). In P. yezoensis, on the other hand, Gao et al. (1992)
found evidences for active HCO3− transport, in that CO2 uptake was extremely
slow compared with the photosynthesis, and that external CA was never found in
5. Porphyra in IMTA
According to the FAO (2009b), in 2006, 47% of the world’s fish food supply
was produced in aquaculture. This activity continues to grow more rapidly than
all other animal food-producing sectors and in 2006 accounted for 36% of the
world total aquatic animal production by weight. In turn, in 2007, aquatic plants
aquaculture represents nearly 23% of the world aquaculture production, based
principally in Asian countries. The seaweeds that are produced are for human
consumption or for extraction of hydrocolloids (agar, alginates or carrageenan).
There are two main biological techniques for treatment of animal aquaculture
waters: bacterial dissimilation into gases and plant assimilation into biomass.
Bacterial biofilters allow effective and significant aquaculture water recirculation
(van Rijn, 1996), but the technology is not simple and such systems usually accumulate nitrate and sludge that need to be disposed, and are expensive to operate.
Biofiltration by algae is assimilative (Krom et al., 1989) and they use excess nutrients (particularly C, N, and P) to produce new biomass that can easily be removed
from the water and that can have an economic value. In China, the annual production of over 7.4 Mt of seaweed (FAO, 2009a) may be responsible for removal
of more than 40,000 t of nitrogen (N) from coastal waters. It is clear that seaweed
production can help mitigate the potential environmental impacts of animal
RUI PEREIRA AND CHARLES YARISH
production and contribute to the development of an environmentally and
economically sustainable aquaculture.
The principle of Integrated Multi-trophic Aquaculture (IMTA) systems is to
build a simplified ecosystem where the resources provided, mainly feed and water,
will be used by two to three other extractive aquaculture organisms: molluscs and
seaweeds. This allows a system to virtually use much of the nutrients minimising
the production of wastes. The best examples of IMTA systems operating in
different parts of the world were described by Chopin et al. (2008).
Because of its morphological characteristics, high surface area to volume
ratio (SA/V), Porphyra is one of the most promising species to be used as biofilter
in IMTA. The thallus of Porphyra is a thin blade with one or two layers of cells,
all potentially involved in nutrient absorption. It can be argued that a thallus with
high surface area to volume ratio does not allow storage of nutrients in reserve
tissues like those of brown algae (e.g. Laminariales and Fucales). The advantage
of Porphyra is its rapid growth, over 25% day−1 (Pereira et al., 2006; Carmona
et al., 2006), which can allow repeated harvests and continuous removal of nutrients from the water. There are yet other factors supporting the use of Porphyra in
IMTA. When compared with most other seaweeds, besides its rapid growth rate,
Porphyra has high nutrient uptake rates, is capable of coping with high NH4+
concentrations and is able to store N in its tissue up to 6% dry weight (dw).
Furthermore, the biomass produced has several valuable applications.
The best growth rates and nutrient removal capacities are usually found in
species with high surface area to volume ratios as explained by the functionalform model (Hanisak et al., 1990; Littler and Littler, 1980). For that reason, a lot
of work has been done using thin blade-like species of Ulva (Chlorophyta).
Martínez-Aragón et al. (2002) compared phosphate removal, from sea bass cultivation effluents, by Ulva rotundata, Ulva intestinalis (formerly Enteromorpha
intestinalis) and Gracilaria gracilis. The maximum P uptake rate (2.86 mmol PO4−3
g−1 dw h−1) was found in U. rotundata. In a follow-up study with the same species
(Hernández et al., 2002), U. rotundata also showed the highest NH4+ uptake rate,
89.0 mmol g−1 dw h−1. Chung et al. (2002) recorded an uptake rate of 114.6 mmol
NH4+ g−1 dw h−1 for U. pertusa. Fujita (1985) obtained, for U. lactuca, uptake rates
between 2 × 103 and 3.6 × 103 mg N g−1 dw h−1. The same author obtained, for
U. intestinalis, a maximal uptake of 14 × 103 mg N g−1 dw h−1. Mata et al. (2006)
reported what is likely to be the highest N uptake rate for a seaweed in integrated
aquaculture environment. In that work, the authors report a total ammonia nitrogen
(TAN) uptake rate of 90 µmol l−1 h−1 at a TAN flux of about 500 µmol l−1 h−1 and
with 5 g fresh weight (fw) l−1 stocking density. This is the equivalent to approximately
4.5 mg N day−1 g−1 fw of the alga.
The results of N uptake by Porphyra, although lower than those reported for
A. armata, are very interesting and promising. In laboratory conditions, Pereira
et al. (2006) obtained an N uptake rate of 1.5 mg N day−1 g fw of P. dioica. Also
in the laboratory, Carmona et al. (2006) obtained a maximum removal of 1.75 mg
N day−1 g−1 fw of P. purpurea and P. haitanensis. In both cases, tissue N recorded
THE ROLE OF PORPHYRA IN SUSTAINABLE CULTURE SYSTEMS
was over 6% dw. As mentioned earlier, Porphyra has other characteristics that
constitute important advantages when choosing a biofilter. Particularly relevant,
in terms of the potential role of Porphyra in ecosystem sustainability, is the ability
of some species to uptake N equally well in the form of NH4+ or NO3− as shown
by Pereira et al. (2006) and Carmona et al. (2006).
The results obtained so far show that Porphyra has an important role in the
nutrient balance in the ecosystems. If we consider a 10:1 dw:fw ratio and average
values of 5.5% N, 0.7% P and 38% C in tissue dw, we realise that the aquaculture
of Porphyra removes, annually, a significant amount of these nutrients from water.
In fact, considering FAO last production figures (FAO, 2009a), approximately
1.51 Mt of Porphyra were produced in 2007. This may equal to a removal of
9,063 t of N, 1,057 t of P and 57,404 t of C, every year.
These results are not just theoretical but supported by field studies of production of Porphyra and its nutrient removal capacity (e.g. Chopin et al., 1999; He
et al., 2008). In the most recent paper, He et al. (2008) showed the bioremediation
capability and efficiency of large-scale Porphyra cultivation in the removal of inorganic nitrogen and phosphorus from open sea area. The study took place in
2002–2004, in a 300 ha nori farm along the Lusi coast, Qidong County, Jiangsu
Province, China. Nutrient concentrations were significantly reduced by the seaweed cultivation. Compared with a control area, Porphyra farming resulted in the
reduction of NH4–N, NO2–N, NO3–N and PO4–P by 50–94%, 42–91%, 21–38%
and 42–67%, respectively. Nitrogen and phosphorus contents in dry Porphyra
thalli harvested from the Lusi coast averaged 6.3% and 1.0%, respectively. The
authors concluded that the annual biomass production of P. yezoensis (about 800
kg dw ha−1 from a 300 ha cultivation) equaled to an average of 14,708 kg of tissue
N and 2,373 kg of tissue P harvested. These results indicate clearly that Porphyra
can efficiently remove excess nutrient from nearshore eutrophic coastal areas.
The nutrient uptake capacity of Porphyra can also be useful in inland IMTA
systems, providing biofiltration for intensive fish aquacultures. The effluents from
these systems can reach high concentrations of both NH4–N and NO3–N.
Porphyra is capable of coping with high concentrations N in either form whether
for uptake or growth.
The main constraint in the case of land-based tank cultivation is the area of
cultivation needed to significantly reduce the nutrient loads of the effluents.
For instance, for an effluent with 143 µM N and 10 µM P at a flux of 190 l min−1,
Carmona et al. (2006) calculates that 282 m2 of Porphyra amplissima cultivation
are needed to remove 90% of the N. Pereira et al. (2006), for Porphyra dioica,
calculated that 179 m2 of culture area would be needed to remove 50% of the N.
This is considering that approximately 600 g of N are released per day by ton of
fish (data from a 50 fish pond, 36 m3 each, turbot and sea bass farm, as described
by Matos et al., 2006). Those authors refer, however, that in an IMTA system, the
performance of P. dioica is likely to improve. Under those tank cultivation conditions, the algae would have more nutrients and CO2 available, owing to the constant
water flux. This would also allow to experiment with higher stocking densities,
RUI PEREIRA AND CHARLES YARISH
Biomass needed (kg)
15 -1 )
Figure 2. Three-dimensional simulation model of the biomass of Porphyra dioica needed to achieve
an 80% reduction in a N load (during 12 h light day) of 600 g N day−1 (195 l min−1 × 150 µM-N).
possibly increasing nutrient removal. Ultimately, the best nutrient removal
performance would be the result of the conditions that could yield the optimal
combination of growth and tissue N content, for a given stocking density (Fig. 2).
It is clear that Porphyra can play an important role in the sustainable development of intensive aquaculture. Species of Porphyra have high ammonia uptake
rates, high ammonia uptake efficiencies, high yields and high protein contents
(N content). Detailed economic studies are now needed to confirm the feasibility
of such systems.
In conclusion, we think Porphyra is a very promising marine organism for
biotechnological exploitation, one of the best seaweed for application in sustainable aquaculture (IMTA) and an interesting organism to be used as a model in
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Biodata of Gamze Turan and Amir Neori, authors of “Intensive Seaweed Aquaculture:
A Potent Solution Against Global Warming”
Dr. Gamze Turan is a faculty member at Ege University, Fisheries Faculty, Aquaculture
Department, Izmir, Turkey. She obtained her Ph.D. from Ege University Institute
of Natural and Applied Sciences in 2007 in Aquaculture and continued her research
on seaweeds, their cultivation, and usage. Dr. Turan’s scientific interests cover the
areas of seaweed cultivation, and their usage as phycocolloids, human food, animal
feed, fish feed, fertilizers and soil conditioners, biomass for fuel, cosmetics, integtated
multitrophic aquaculture (IMTA), wastewater treatment, etc. She has published
over 20 peer-reviewed publications.
Dr. Amir Neori is a senior scientist at the Israel Oceanographic & Limnological
Research, Ltd. The National Center for Mariculture, Eilat, Israel. He obtained
his Ph.D. from the University of California San Diego – Scripps Institution of
Oceanography in 1986 in Marine Biology and continued his research in sustainable
mariculture and algae in his present capacity. Dr. Neori’s scientific interests are
in the area of environmentally-friendly aquaculture, algal aquaculture, reduction in
aquaculture environmental impact, integrated multitrophic aquaculture (IMTA), and
biofuel from algae. He has published over 70 peer-reviewed publications.
E-mail: email@example.com; firstname.lastname@example.org
A. Israel et al. (eds.), Seaweeds and their Role in Globally Changing Environments,
Cellular Origin, Life in Extreme Habitats and Astrobiology 15, 357–372
DOI 10.1007/978-90-481-8569-6_20, © Springer Science+Business Media B.V. 2010
INTENSIVE SEA WEED AQUACULTURE: A POTENT SOLUTION
AGAINST GLOBAL WARMING
GAMZE TURAN1 AND AMIR NEORI2
Fisheries Faculty, Aquaculture Department, Ege University,
Bornova, Izmir, 35100, Turkey.
Israel Oceanographic & Limnological Research Ltd, National Center
for Mariculture, P.O. Box 1212, Eilat, 88112, Israel
On the basis of current understanding of the relationship between climate change
and energy policy, development of an effective and multistructured renewable energy
sector is crucial, as acknowledged in the United Nations Framework Convention on
Climate Change (UNFCCC) and the fourteenth Conference of the Parties (COP14), held in December 2008 in Ponzan, Poland. The worldwide energy demand is
increasing rapidly as many industries and populations are rapidly expanding. Since
fossil fuels are finite resources and their combustion leads to a further increase of
greenhouse gases, such as CO2, SO2, and NOx, their continued use is not sustainable.
Today, renewable energy sources supply 14% of the total global energy demand.
Some expect that in 2040, 50% of the world energy supply will come from renewable
sources (Demirbas, 2008). Additional efforts and further research and development
on biofuels, toward environmentally and economically sustainable processes, are
essential for the full exploitation of this given market opportunity.
The substitution of conventional fuels by biofuels can reduce pollution and
support sustainability. First-generation biofuels, such as biodiesel and bioethanol
derived from biomass, have their environmental benefits related to carbon-neutral
energy. However, increasing biofuel production from land crops strains the global
food supply. Owing to these limitations, second-generation (bio) fuels – from
biomasses that generate carbon neutral energy without competing with food production – have been developed. These can be produced from the residual nonfood
parts of current crops, as well as novel energy crops, such as seaweeds.
The culture of seaweeds has unique characteristics, which make it different
and in many ways attractive in comparison with other biofuel sources. Seaweeds,
also known as “marine macroalgae,” “aquatic plants,” or “sea vegetables,” are
autotrophic organisms that produce biomass using sunlight and extracting from
the water-dissolved inorganic nutrients, including carbon. Several seaweed species