CO2 enrichment and plant response, change in quality and quantity of crop produce
The beneficial effects of atmospheric CO2 enrichment may be divided into three distinct
growth response phases. First is a well-watered optimum-growth-rate phase where
a 300 parts per million increase in the CO2 content of
the air generally increases plant productivity by approximately 30%.
Next comes a nonlethal water-stressed phase where the same
increase in atmospheric CO2 is more
than half again as effective in increasing plant productivity. Finally, there
is a water-stressed phase normally indicative of impending death, where
atmospheric CO2 enrichment may actually
prevent plants from succumbing to the rigors of the environment and enable them
to maintain essential life processes, as life ebbs from corresponding
ambient-treatment plants.
Empirical records provide incontestable evidence of global changes:
foremost among these changes is the rising concentration of CO2 in
the earth's atmosphere. Plant growth is nearly always stimulated by elevation
of CO2. Photosynthesis increases, more plant biomass accumulates per
unit of water consumed, and economic yield is enhanced.
The profitable use of
supplemental CO2 over years of greenhouse practice points to
the value of CO2 for plant production. Plant responses to CO2 are
known to interact with other environmental factors, e.g. light, temperature, soil
water, and humidity. Important stresses including drought, temperature,
salinity, and air pollution have been shown to be ameliorated when CO2 levels
are elevated. In the agricultural context, the growing season has been
shortened for some crops with the application of more CO2; less
water use has generally, but not always, been observed and is under further
study; experimental studies have shown that economic yield for most crops
increases by about 33% for a doubling of ambient CO2 concentration.
However, there are some reports of negligible or negative effects.
Plant species respond differently to CO2 enrichment,
therefore, clearly competitive shifts within natural communities could occur.
Thought of less importance in managed agro-ecosystems, competition between
crops and weeds could also be altered. Tissue composition can vary as CO2 increases
(e.g. higher C: N ratios) leading to changes in herbivory, but tests of crop
products (consumed by man) from elevated CO2 experiments have
generally not revealed significant differences in their quality. However, any
CO2-induced change in plant chemical or structural make-up could
lead to alterations in the plant's interaction with any number of environmental
factors—physicochemical or biological. Host-pathogen relationships, defense
against physical stressors, and the capacity to overcome resource shortages
could be impacted by rises in CO2.
Root biomass is known
to increase but, with few exceptions, detailed studies of root growth and
function are lacking. Potential enhancement of root growth could translate into
greater rhizodeposition, which, in turn, could lead to shifts in the
rhizosphere itself. Some of the direct effects of CO2 on
vegetation have been reasonably well-studied, but for others work has been
inadequate. Among these neglected areas are plant roots and the rhizosphere.
Therefore, experiments on root and rhizosphere response in plants grown in CO2-enriched
atmospheres will be reviewed and, where possible, collectively integrated.
In general, C3 plant
species are more responsive to atmospheric carbon dioxide (CO2)
enrichment than C4-plants. Increased relative growth rate at
elevated CO2 primarily relates to increased Net Assimilation
Rate (NAR), and enhancement of net photosynthesis and reduced photorespiration.
Transpiration and stomatal conductance decrease with elevated CO2,
water use efficiency and shoot water potential increase, particularly in plants
grown at high soil salinity. Leaf area per plant and leaf area per leaf may
increase in an early growth stage with increased CO2, after a period
of time
Leaf Area Ratio (LAR) and Specific Leaf Area (SLA) generally decrease.
Starch may accumulate with time in leaves grown at elevated CO2. Plants
grown under salt stress with increased (dark) respiration as a sink for
photosynthates, may not show such acclimation to increased atmospheric CO2 levels.
Plant growth may be stimulated by atmospheric carbon dioxide enrichment
and reduced by enhanced UV-B radiation but the limited data available on the
effect of combined elevated CO2 and ultraviolet B (280–320 nm)
(UV-B) radiation allow no general conclusion. CO2-induced increase
of growth rate can be markedly modified at elevated UV-B radiation.
Plant responses to elevated
atmospheric CO2 and other environmental factors such as soil
salinity and UV-B tend to be species-specific, because plant species differ in
sensitivity to salinity and UV-B radiation, as well as to other environmental
stress factors (drought, nutrient deficiency). Therefore, the effects of joint
elevated atmospheric CO2 and increased soil salinity or
elevated CO2 and enhanced UV-B to plants are physiologically
complex.
High CO2
Makes Crops Less Nutritious
Crops grown in the high-CO2
atmosphere of the future could be significantly less nutritious. a new
challenge as society reckons with both rising carbon emissions and malnutrition
in the future.
Scientists generally predict that crop yields could
fall in a warmer world—though higher atmospheric CO2 by itself
should raise yields, as plants find it easier to extract CO2 from
the air to make carbohydrates.
The effect climate change might have on the nutritional
value of crops, as opposed to their yield, has been even murkier. Previous
studies have given conflicting results.
The CO2 levels expected in the second half of
this century will likely reduce the levels of zinc, iron, and protein in wheat,
rice, peas, and soybeans. Some two billion people, the researchers note, live
in countries where citizens receive more than 60 percent of their zinc or iron
from these types of crops. Deficiencies of these nutrients already cause an
estimated loss of 63 million life-years annually.
C3 Crops Hit Hardest
Conducted over six growth years on field sites in Japan,
Australia, and the United States, the study compared crops grown in normal
conditions with ones grown in nearby experimental plots where the air is
enriched with CO2 via open-air sprayers. The current atmospheric CO2
level is 400 parts per million; in the enriched plots, it was between 546
and 586 parts per million, a level scientists expect the atmosphere to reach in
four to six decades.
In addition to wheat, rice, peas, and soybeans, which all
use a form of photosynthesis known as C3, corn and sorghum, which
use C4 photosynthesis, a faster kind. There was relatively little
effect of CO2 enrichment on the nutritional value of the C4
crops.
In the C3 crops, however, there was
significant declines in zinc and iron. The largest was a 9.3 percent drop in
the zinc level in wheat. They also found reduced levels of protein in wheat,
rice, and peas, but not in soybeans. In general, crops are losing nutrients as
CO2is going up.
Unfortunately, the new study sheds little light on why
more CO2 in the atmosphere should mean less nutritious plants. One
hypothesis has been that plants in an enriched atmosphere produce so much
carbohydrate that it dilutes the other nutrients.
The new study seems to rule out that hypothesis: Instead
of a uniform dilution of all other nutrients in the crops, it found that
nutrients changed unevenly when CO2 was higher.
Quality and Quantity
The need to balance changes in yield against changes in
the nutritional value of crops makes predicting the future of agriculture an
even more complicated task.
"Rising global CO2 increases yield and
decreases water use by crops, and this is often presented as one positive of
atmospheric change, environments will mean less nutritional crops, so that
"increased quantity is at the expense of quality."
There should be a global effort to develop new breeds of
wheat, rice, peas, and soybeans that show resistance to higher CO2
levels. While the various cultivars of wheat, peas, and soybeans in the study
all suffered similar nutrient losses in response to higher CO2, rice
offered a ray of hope: Its cultivars varied wildly in their response.
CO2 concentration
CO2 is
vital for plant growth, as it is the substrate for photosynthesis. Plants take
in CO2 through stomatal pores on their leaves. At the
same time as CO2 enters the stomata, moisture escapes. This
trade-off between CO2 gain and water loss is central to plant
productivity. The trade-off is all the more critical as Rubisco, the
enzyme used to capture CO2, is efficient only when there is a high
concentration of CO2 in the leaf. Some plants overcome this
difficulty by concentrating CO2 within their leaves
using C4 carbon fixation or Crassulacean acid
metabolism. However, most species used C3 carbon
fixation and must open their stomata to take in CO2 whenever
photosynthesis is taking place.
The
concentration of CO2 in the atmosphere is rising due
to deforestation and the combustion of fossil fuels. This would
be expected to increase the efficiency of photosynthesis and possibly increase
the overall rate of plant growth. This possibility has attracted considerable
interest in recent years, as an increased rate of plant growth could absorb
some of the excess CO2 and reduce the rate of global
warming. Extensive experiments growing plants under elevated CO2 using Free
Air Concentration Enrichment (FACE) have shown that photosynthetic efficiency
does indeed increase. Plant growth rates also increase, by an average of 17%
for above-ground tissue and 30% for below-ground tissue. However, detrimental
impacts of global warming, such as increased instances of heat and drought
stress, mean that the overall effect is likely to be a reduction in plant
productivity.
Reduced
plant productivity would be expected to accelerate the rate of global warming.
Overall, these observations point to the importance of avoiding further
increases in atmospheric CO2 rather than risking runaway
climate change.
A regional climate change model (PRECIS) for China, developed by the
UK's Hadley Centre, was used to simulate China's climate and to develop climate
change scenarios for the country. Results from this project suggest that,
depending on the level of future emissions, the average annual temperature
increase in China by the end of the twenty-first century may be between 3 and 4 °C. Regional crop models were driven by PRECIS output to
predict changes in yields of key Chinese food crops: rice, maize and wheat.
Modelling suggests that climate change without carbon dioxide (CO2)
fertilization could reduce the rice, maize and wheat yields by up to 37% in the
next 20–80 years. Interactions of CO2 with limiting factors,
especially water and nitrogen, are increasingly well understood and capable of
strongly modulating observed growth responses in crops.
More complete reporting of free-air
carbon enrichment experiments than was possible in the Intergovernmental Panel
on Climate Change's Third Assessment Report confirms that CO2 enrichment
under field conditions consistently increases biomass and yields in the range
of 5–15%, with CO2 concentration elevated to 550 ppm Levels of CO2 that are elevated to more
than 450 ppm will probably cause some deleterious effects in grain
quality. It seems likely that the extent of the CO2 fertilization
effect will depend upon other factors such as optimum breeding, irrigation and
nutrient applications.
The modelling work
took into account climatic variables, irrigation, soil variables and the
influence of higher atmospheric concentrations of carbon dioxide (CO2)
on plant metabolism. In general, climate change itself tends to reduce crop
yield but the fertilization effect of CO2 tends to increase
yield. The balance between these two effects is likely to depend, in reality,
on factors such as the availability of water and nutrients and the prevalence
of pests and diseases, all of which are also likely to be affected by climate
change.
CO2 fertilization
effectively offsets yield decreases caused by shorter growth duration due to
higher temperatures in rice
If the direct
effect of CO2 is included, average yields are projected to
increase for rainfed maize and decrease for irrigated maize in the 2080s. The
increase is likely to be highest for rainfed maize because the higher CO2 concentration
would boost the yield of rainfed maize under the current water-limited
conditions prevalent in North China (the biggest maize cultivation area).
Without the CO2 fertilization effect, the average yield of both
rainfed and irrigated maize is likely to fall because the higher temperature
may shorten the growth period by between 4 and 8 days. While irrigation might
counteract the trend towards a decrease in yield (assuming sufficient water is
available), it is not expected to stop it completely.
Yield decreases
would be greatest if higher temperatures occur during the period when the maize
ears are swelling. These results show a large relative benefit to maize yields
from elevated CO2. This is in contrast to most C4 crop
experiments which show minor absolute changes in yield due to CO2 enrichment.
As with rice, there is large regional variability in the yield change. This
could be due to the use of calibrated irrigation and nutrition parameters in
the model which were validated under present CO2 concentration
rather than in a higher CO2 environment.
In wheat, if
the effect of CO2 fertilization is included, average wheat
yields are shown to increase in China in the 2020s, 2050s and 2080s for both
rainfed and irrigated wheat. Spatial variability is again large. The response
of wheat to future atmospheric CO2 increases is likely to
significantly constrain potential increases in yield. But for irrigated wheat
to benefit from the effects of CO2 fertilization, nutrients
need to be non-limiting. Without CO2 fertilization, wheat yields are
expected to be 10-20% lower by 2080 compared with current yields.
Effects
of elevated CO2 on grain quality
Wheat
of two genotypes was grown in the field under CO2 gradient
enrichment (CGE—half of open) with a controlled chamber in the Chinese Academy
of Agricultural Science experiment station in Beijing in 2001–2002. The
gradient CO2 enrichment was from 451 to 565 mg kg−1. Measurement
for effects of elevated CO2 on grain quality showed that: (i)
protein content for flour was found to significantly decrease with CO2concentration
gradient enrichment (at range 57 μmol
mol−1; (ii)the
sedimentation value of ZhongYu five was found to decrease a little. Even though
there were some errors and uncertainties due to limited samples, significant
differences between different varieties still exist after strict measurements.
These results indicate that elevated CO2 levels may cause a
decrease in the quality of bread wheat due to generally lowered protein
content.
The
above results from CGE experiments have confirmed the results from previous
studies: elevated CO2 can cause more or less deleterious
effects on grain quality. In addition, there would be distinct differences
between varieties. Most of the experiments showed reductions in grain nitrogen
content or grain protein at elevated levels of CO2, although some
found no significant effect of elevated CO2 on grain quality.
Rises
in the concentration of CO2 in the atmosphere are likely to be
accompanied by temperature increases. Small increases in temperature (2–4 °C) had a larger effect than
elevated CO2 on grain quality. Moreover, the effects of
elevated CO2 on grain quality may be partially balanced because
temperature increases can enhance grain protein content. However, it is
unlikely that any high temperature effects will totally compensate for CO2 enrichment.
Data from Kimball's experiments (2001) suggest that adequate fertilizer is
necessary to attain good quality grain and that with ample fertilizer the
deleterious effects of elevated CO2 will be minor. Furthermore,
crops grown with limiting levels of nitrogen probably have poorer quality grain
than they could have. CO2 enrichment in the atmosphere during
coming decades is likely to make the quality poorer still.
Potential
adaptation technologies of acclimation to CO2 fertilization
Measurements
have shown that with prolonged exposure to elevated atmospheric CO2,
the photosynthetic rate gradually declined, approaching or even less than that
in ambient. These results indicate an acclimation or down regulation to the
higher CO2 levels. But CO2 fertilization still can
be favoured for adaptation in a future climate. A detailed understanding of CO2 fertilization
should be taken into account in developing adaptation technology.
New
varieties by seed selection are one of the key methods of increasing crop yield
and improving crop quality as well as adapting to environment change. With
increasing ambient CO2 concentration and a warmer climate,
especially in winter, new crop varieties with high yield, warm-winter
resistance under higher CO2 should be favoured for adaptation
in a future climate. For rice, cross breeding of Indica and Japanica varieties
is considered ideal for enhancing morphological characteristics.
Improving
crop cultivation would be another helpful technique for acclimation of crops to
the CO2 fertilization effect. For example, adjusting crop
planting time could avoid light energy loss while adjusting the planting area
and region of C3 and C4 crops and increasing plant
density could increase the accumulation and efficient use of CO2.
It
is very difficult to understand the interactive impact of elevated atmosphere
CO2 and raising temperature on crop growth and yield formation.
More CO2 fertilization can be practiced through adjusting
planted crop distributions and sowing times.
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