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Ethiopia goes organic to feed herself - The Institute of Science In ...

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39<br />

Woody trees decompose slowly and are an important carbon sink<br />

main types <strong>of</strong> lignin, guaiacyl lignin<br />

and guaiacyl-syringyl. Guaiacyl<br />

lignin is characteristic <strong>of</strong> s<strong>of</strong>twoods,<br />

which are resistant <strong>to</strong><br />

chemical and biological degradation.<br />

Guaiacyl-syringyl lignin is typical<br />

<strong>of</strong> hardwoods such as poplar,<br />

which are more readily degraded.<br />

Modifying plants with a gene<br />

enhancing the proportion <strong>of</strong> guaiacyl-syringyl<br />

lignin therefore provides<br />

a lignin more readily degraded<br />

by chemicals or enzymes.<br />

Reducing lignin content also leads<br />

<strong>to</strong> plants more readily digested with<br />

enzymes or chemicals.<br />

Lignin reduction has been<br />

achieved using anti-sense genes <strong>to</strong><br />

limit production <strong>of</strong> key enzymes on<br />

the lignin biosynthesis pathway.<br />

Multiple genetic transformations <strong>of</strong><br />

forest trees have been used <strong>to</strong><br />

enhance production <strong>of</strong> guaiacylsyringyl<br />

lignin and <strong>to</strong> limit <strong>to</strong>tal<br />

lignin production. Four<br />

Agrobacterium T-DNA vec<strong>to</strong>rs,<br />

each with a cauliflower mosaic<br />

virus promoter, two <strong>of</strong> which included<br />

anti-sense <strong>to</strong> limit undesirable<br />

enzymes and two with sense constructions<br />

<strong>to</strong> enhance desirable<br />

enzymes, were used <strong>to</strong> simultaneously<br />

alter the genome <strong>of</strong> aspen<br />

(Populus tremuloides). This resulted<br />

in reduced lignin content <strong>of</strong> guaiacyl<br />

lignin and increased guiaicylsyringyl<br />

proportion in the remaining<br />

lignin.<br />

Even though a potentially desirable<br />

end product is created, the<br />

multiple transformations (gene<br />

stacking) are liable <strong>to</strong> create chromosome<br />

instability leading <strong>to</strong><br />

translocations, duplications and<br />

deletions through homologous<br />

recombination during germ cell formation<br />

and in somatic tissues<br />

(mi<strong>to</strong>tic<br />

recombination).<br />

<strong>In</strong>dependent studies <strong>of</strong> transgene<br />

integration using T-DNA vec<strong>to</strong>rs in<br />

aspen showed extensive DNA<br />

sequence scrambling at the insertion<br />

points. DNA sequence scrambling<br />

occurring in the cells during<br />

growth is a significant complication<br />

in long-lived trees.<br />

Lignin genetic engineering is<br />

promoted as a promising strategy<br />

<strong>to</strong> improve fibre production but the<br />

drawbacks <strong>of</strong> anti-sense manipulation<br />

and transgene stability are not<br />

seriously dealt with. Trees genetically<br />

modified <strong>to</strong> produce low lignin<br />

are called "super" trees with little<br />

consideration <strong>of</strong> pest resistance<br />

and genetic stability. Field and<br />

pulping performance <strong>of</strong> transgenic<br />

poplars with altered lignin was<br />

evaluated <strong>to</strong> be superior by the<br />

developers <strong>of</strong> the poplar and abnormal<br />

pest damage was not found.<br />

However, the pest damage studies<br />

were cursory and not compared<br />

with experimental controls, but with<br />

norms reported by government<br />

agencies.<br />

<strong>The</strong> antibiotic resistance markers<br />

from the leaves <strong>of</strong> transgenic<br />

aspen have been studied for their<br />

persistence in the soil. <strong>The</strong> field<br />

study showed that the marker DNA<br />

<strong>of</strong> the aspen leaves persisted for as<br />

much as four months in the soil.<br />

<strong>The</strong> persistence <strong>of</strong> antibiotic resistance<br />

genes in the forest ecosystem<br />

is likely <strong>to</strong> impact not only soil<br />

microbes, but human and animal<br />

inhabitants <strong>of</strong> the forest as well.<br />

Lignin content increases as<br />

crops age or are stressed. Animal<br />

<strong>feed</strong> rich in lignin is poorly<br />

digestible and considered <strong>to</strong> be <strong>of</strong><br />

low quality. Grass, alfalfa or maize<br />

with reduced lignin or lignin with<br />

increased guaiacyl-syringyl proportion<br />

(readily digested) may provide<br />

a large economic benefit in animal<br />

production, provided that the<br />

genetic modifications do not result<br />

in susceptibility <strong>to</strong> preda<strong>to</strong>ry<br />

insects, fungi and bacteria and do<br />

not compromise food or <strong>feed</strong> safety<br />

(for example, fungus food contamination<br />

may lead <strong>to</strong> pollution <strong>of</strong> food<br />

with <strong>to</strong>xins, causing liver damage<br />

and cancer).<br />

<strong>The</strong> main technique used <strong>to</strong> produce<br />

lignin modifications is antisense<br />

genes designed <strong>to</strong> reduce<br />

one or another enzyme level on the<br />

pathway <strong>to</strong> lignin production. Maize<br />

with improved forage quality was<br />

produced by down-regulating the<br />

enzyme O-methyl transferase <strong>to</strong><br />

limit lignin production. Tall fescue<br />

pasture grass with improved forage<br />

digestibility was produced using an<br />

anti-sense gene for the lignin precursor<br />

enzyme cinnamyl alcohol<br />

dehydrogenase. Alfalfa down-regulated<br />

for lignin enzyme caffeoyl<br />

coenzyme A 3-O-methyl transferase<br />

produced plants with<br />

increased guaiacyl-syringyl lignin<br />

proportions leading <strong>to</strong> improved<br />

rumen digestibility.<br />

<strong>The</strong>re is little question that the<br />

forage and fodder with reduced<br />

lignin and lignin with improved<br />

composition are more desirable<br />

food sources for grazing animals.<br />

However, the downside <strong>of</strong> lignin<br />

manipulation - greater disease susceptibility<br />

- was not thoroughly considered<br />

by developers <strong>of</strong> crops with<br />

modified lignin. <strong>The</strong> developers<br />

seem <strong>to</strong> ignore safety issues while<br />

they promote the modified crops.<br />

Furthermore, smooth brome<br />

grass clones selected using conventional<br />

breeding showed that<br />

reduced lignin was associated with<br />

severe rust fungus disease. Alfalfa<br />

selected for forage quality (including<br />

reduced lignin) had reduced<br />

vigour but was not expected <strong>to</strong><br />

affect levels <strong>of</strong> disease resistance.<br />

Sudan grass selected for brownmidrib<br />

trait (an indica<strong>to</strong>r <strong>of</strong> reduced<br />

lignin) experienced severe yield<br />

reductions and environmental sensitivity,<br />

particularly during cooler<br />

growing seasons.<br />

Lignin modification <strong>of</strong> trees and<br />

forage crops has been a focus <strong>of</strong><br />

research in genetic engineering.<br />

But lignin provides both fundamental<br />

structural features and resistance<br />

<strong>to</strong> animal and microbial pests.<br />

Lignin enhancement that leads <strong>to</strong><br />

greater forage or tree pulp quality<br />

also leads <strong>to</strong> susceptibility <strong>to</strong> disease,<br />

while lignin enhancement<br />

that leads <strong>to</strong> greater disease resistance<br />

makes forage less digestible<br />

and tree pulp more expensive <strong>to</strong><br />

process.<br />

<strong>The</strong> economic consequences <strong>of</strong><br />

effective lignin modification could<br />

be tremendous, but producing<br />

forests and rangelands highly susceptible<br />

<strong>to</strong> insects, fungi and bacteria<br />

would lead <strong>to</strong> economic and<br />

environmental disaster. <strong>The</strong> low<br />

lignin trait is comparable <strong>to</strong> a loss<br />

in immune functions comparable <strong>to</strong><br />

AIDS in mammals. <strong>The</strong> chemical<br />

corporations might well welcome a<br />

huge increase in pesticides <strong>to</strong> fight<br />

disease in forests and pastures.<br />

Nevertheless, the best strategy is<br />

<strong>to</strong> proceed prudently and honestly<br />

evaluate the consequences <strong>of</strong> far<br />

reaching genetic engineering<br />

experiments.<br />

Note added by edi<strong>to</strong>r: Another<br />

consideration is ecological. Wood,<br />

with its naturally high lignin content,<br />

generally takes a long time <strong>to</strong><br />

decay and recycle in the ecosystem,<br />

probably for good reasons. It<br />

is a long-term energy s<strong>to</strong>re complementing<br />

the shorter-term energy<br />

s<strong>to</strong>rage depots, which enables the<br />

ecosystem <strong>to</strong> function most efficiently<br />

and effectively (see "Why<br />

are organisms so complex? A lesson<br />

in sustainability", SiS 21).<br />

Slow-decaying wood is also a major<br />

carbon sink. Reducing its lignin<br />

content <strong>to</strong> enhance degradation will<br />

end up returning carbon dioxide <strong>to</strong>o<br />

rapidly <strong>to</strong> the atmosphere, thereby<br />

exacerbating climate change (see<br />

"Why Gaia needs rainforests" SiS<br />

20). SiS<br />

www.i-sis.org.uk

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