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Svirski et al. 1993 ), and the extent of the damage is clearly determined by the
intensity of the infections (Cancino et al. 1987 ; Buschmann and Gomez 1993 ).
Farming of Gracilaria chilensis during the last 40 years is a clear example of how
detrimental epiphytes may become (Kuschel and Buschmann 1991 ; Pizarro and
Santelices 1993 ).
In addition, negative effects on the economic value of brown and red algae infested
by endophytes have been reported (Yoshida and Akiyama 1979 ;Apt 1988a , b ). The
endophytic phaeophycean Laminariocolax ( Streblonema) aecidioides causes tissue
thickening in the commercially valuable alga Undaria sp. (Yoshida and Akiyama
1979 ), and Streblonema -like endophytes are known to produce galls in some algal
hosts (Andrews 1977 ;Apt 1988a ).
Estimates of economic losses for Japan and Korea (Gachon et al. 2010 ) are
estimated at about 10% of the annual production (0.15 billion US dollars globally),
mostly due to the oomycete pathogens Olpidiopsis spp . and Pythium spp. (Ding
and Ma 2005 ; Park et al. 2000 ) . However, spatial and interannual variability
feature dramatic variations with reductions of 15-20% and 20-30% in yields and
value, respectively, in South Korea and losses in Japan rose to 64% in 1993
(Gachon et al. 2010 ).
Removal of epiphytes can be carried out by physical (mechanical brushing, rapid
water movement), chemical (rinsing in chlorine, copper, or changes in pH), and
biological means (use of epiphyte herbivores). However, type III to type V epiphytes
(see above), as well as endophytes and eukaryotic parasites, cannot be controlled by
such treatments and alternative methods based on the current understanding of the
genetic and biochemical basis of close associations are required in order to limit their
impacts on algal mariculture. Interestingly, a recent study showed that dipping of
seedlings in a commercial extract from Ascophyllum nodosum (AMPEP), prior to
out-planting, was efficient for both improving the daily growth rate and increased
productivity of both varieties of Kappaphycus and could be used as part of a
management protocol to control or reduce the impact of Neosiphonia infection in
commercial cultivation areas in the sea (Borlongan et al. 2011 ).
11.4.3 Fouling
The negative effects of intimate associations on macroalgal fitness can be expected
to exert a strong pressure of selection for efficient antifouling defenses. Chemical
resistance against various epibiotic organisms, e.g., bacteria, microalgae, periphy-
ton, fungi, invertebrate larvae, and epiphytic macroalgae, has been tested for all
major macroalgal groups (worth 1.5 billion US dollars globally reviewed in Davis
et al. 1989 ; Steinberg and deNys 2002 ; Lane and Kubanek 2008 ).
Although a wealth of examples on antifouling properties of algal extracts exists,
the generality of antifouling metabolites has been questioned. The problem with
most of the existing studies is that, although they clearly show macroalgae com-
monly containing metabolites with antifouling properties, it is not known whether
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