Metal hyperaccumulation in plants

During million years the evolution leaded plants to develop different strategies to defence from natural enemies, giving rise to an evolutionary weaponry war in which the survival of ones and others depends into the ability to beat the other’s adaptations. It is in that scenario where the high-level accumulation of heavy metals in plants plays an important role.

INTRODUCTION

Boyd (2012) commented that plant defences can be grouped in different categories:

• mechanic: thorns, coverage, etc.
• chemical: different organic and inorganic components.
• visual: crypsis and mimicry .
• behavioural: related with phenology’s modification.
• and associative: symbiosis with other organisms, such is the case of the genus Cecropia, which has stablished a symbiotic relationship with ants of the genus Azteca, who protects these plants – to know more: Plants and animals can also live in marriage-.

Mechanic defence with thorns (Author: Karyn Christner, Flickr, CC).

It is known that chemical defence is ubiquitous, and thus, a lot of interactions among organisms can be explained for this reason. In this sense, some plants contains high levels of certain chemical elements, frequently metals or metallic components, which plays an important role in the defence, these plants are the heavy metal hyperaccumulating plants.

Heavy metal hyperaccumulating plants and their main characteristics

This plants belong to several families, thus hyperaccumulation is an independent acquisition occurring different times during the evolution. In all cases, hyperaccumulation allowed the ability to grow soils with high levels of heavy metals and to accumulate extraordinary amounts of heavy metals in aerial organs. It is known that the concentration of these chemical elements in hyperaccumulating plants can be 100 – 1000 times higher than in non-hypperaccumulating plants.

Generally, chemistry describes heavy metal as transition metals with atomic mass higher than 20 and with a relative density around 5.  But, from a biological point of view, heavy metals or metalloids are elements which can be toxic in a low concentration. Even though, hyperaccumulating plants has become tolerant, i.e., they hypperacumulate this heavy metals without presenting phytotoxic effects (damage in plant tissues due toxicity).

In this sense, there are three main characteristics typically present in all hyperaccumulating plants:

• Increased absorption rate of heavy metals.
• Roots that perform translocation more quickly.
• Great ability to detoxify and accumulate heavy metals in sheets.

Thus, hyperaccumulating plants are prepared to assimilate, translocate and accumulate high-levels of heavy metals in vacuoles or cellular wall. In part, it is due to the overexpression of genes codifying for membrane transporters.

The threshold values that allow to differentiate a hyperaccumulating plant from a non-hyperaccumulating one are related to the specific phytotoxicity of each heavy metal. According to this criterion, hyperaccumulating plants are plants that when grown on natural soils accumulate in the aerial parts (in grams of dry weight):

• > 10 mg·g^-1 (1%) of Mn or Zn,
• > 1 mg·g^-1 (0,1%) of As, Co, Cr, Cu, Ni, Pb, Sb, Se or Ti
• or > 0,1 mg·g^-1 (0,01%) of Cd.

Minuartia verna, copper hyperacumulating plant (Autor: Candiru, Flickr, CC).

THE ORIGIN OF HYPERACCULATING PLANTS AND THEIR IMPLICATIONS

Till the moment, several hypothesis has been proposed to explain why certain plants can hyperaccumulate heavy metals:

• Tolerance and presence of metals in soils.
• Resistance to drought.
• Interference with other neighbouring plants.
• Defence against natural enemies.

The most supported hypothesis is “Elemental defence”, which indicates that certain heavy metals could have a defensive role against natural enemies, such as herbivores and pathogens. So, in the case these organisms consume plants, they should present toxic effects, which would lead them to die or at least to reduce the intake of this plant in future. Even though heavy metals can act through their toxicity, this does not guarantee plants will not be damaged or attacked before the natural enemy is affected by them. For this reason, it is still necessary a more effective defence which allow to avoid the attack.

In contrast, according to a more modern hypothesis, the “Joint effects”, heavy metals could act along with other defensive organic components giving rise to a higher global defence. The advantages of inorganic elements, including heavy metals, are that they are not synthetized by plants, they are absorbed directly from the soil and thus a lower energetic cost is invested in defence, and also they cannot be biodegraded. Even though, some natural enemies can even avoid heavy metal effects by performing the chelation, i.e., using chelators (substances capable of binding with heavy metals to reduce their toxicity) or accumulating them in organs where their activity would be reduced. This modern hypothesis would justify the simultaneous presence of several heavy metals and defensive organic components in the same plant, with the aim to get a higher defence able to affect distinct natural enemies, which would be expected to do not be able to tolerate different element toxicity.

Thlaspi caerulescens, zinc hyperaccumulating plant (Autor: Randi Hausken, Flickr, CC).

On the other hand, it has been shown that certain herbivores have the ability to avoid the intake of plants with high levels of heavy metals, doing what is called “taste for metals“. Although this is known to occur, the exact mechanism of this alert and avoidance process is still uncertain.

Solanum nigrum, cadmium hyperaccumulating plant (Autor: John Tann, Flickr, CC).

Additionaly, even tough heavy metal concentration in plant are really high, some herbivores manage to surpass this defense by being tolerant, i.e., their diet allows them to intake high dosis of metals and, thus, consume the plant. This could lead to think some herbivores could become specialist in the intake of hyperaccumulating plants, and, thus, this type of defence would be reduced to organisms with varied diets, which are called generalists. It has been demonstrated to not be true, as generalists herbivores sometimes present a higher preference and tolerance for hyperaccumulating plants than specialist organisms.

For all these reasons, it can be said that evolution is still playing an important role in this wonderful weaponry war.

 REFERENCES

• Boyd, R., Davis, M.A., Wall, M.A. & Balkwill K. (2002). Nickel defends the South African hyperaccumulator Senecio coronatus (Asteraceae) against Helix aspersa (Mollusca: Pulmonidae). Chemoecology 12, p. 91–97.
• Boyd, R. (2007). The defense hypothesis of elemental hyperaccumulation: status, challenges and new directions. Plant soil 293, p. 153-176.
• Boyd, R. (2012). Elemental Defenses of Plants by Metals. Nature Education Knowledge 3 (10), p. 57.
• Laskowski, R. & Hopkin, S.P. (1996). Effect of Zn, Cu, Pb and Cd on Fitness in Snails (Helix aspersa). Ecotoxicology and environmentak safety 34, p. 59-69.
• Marschner, P. (2012). Mineral Nutrition of Higher Plants (3). Chennai: Academic Press.
• Noret, N., Meerts, P., Tolrà, R., Poschenrieder, C., Barceló, J. & Escarre, J. (2005). Palatability of Thlaspi caerulescens for snails: influence of zinc and glucosinolates. New Phytologist 165, p. 763-772.
• Prasad, A.K.V.S.K. & Saradhi P.P. (1994).Effect of zinc on free radicals and proline in Brassica and Cajanus. Phytochemistry 39, p. 45-47.
• Rascio, N. & Navari-Izzo, F. (2011). Heavy metal hyperaccumulating plants: How and why do they do it? And what makes them so interesting?. Plant Science 180 (2),p. 169-181.
• Shiojiri, K., Takabayashi, J., Yano, S. & Takafuji, A. (2000) Herbivore-species-specific interactions between crucifer plants and parasitic wasps (Hymenoptera: Braconidae) that are mediated by infochemicals present in areas damaged by herbivores. Applied Entomology and Zoology 35, p. 519–524.
• Solanki, R. & Dhankhar, R. (2011). Biochemical changes and adaptive strategies of plants under heavy metal stress. Biologia 66 (2), p. 195-204.
• Verbruggen, N., Hermans, C. & Schat, H. (2009). Molecular mechanisms of metal hyperaccumulation in plants. New Phytologist 181 (4), p. 759–776.
• Wenzel, W.W. & Jockwer F. (1999). Accumulation of heavy metals in plants grown on mineralised soils of the Austrian Alps. Environmental pollution 104, p. 145-155.

The plants and the climate change

Since a few years ago, we have heard about the climate change. Nowadays, it is already evident and also a concern. This not only affects to us, the humans, but to all kind of life. It has been talked enough about the global warming, but perhaps, what happens to the vegetation has not been much diffused. There are many things affected by climate change and vegetation is also one of them. In addition, the changes in this also affect us. But, what are these changes? how can the vegetation regulate them? And how we can help to mitigate them through plants?

CHANGES ON PLANTS

Biomes distribution

In general, due to climate change, an increase of precipitations in some parts of the world are expected, while in others a decrease is awaited. A global temperature increment is also denoted. This leads to an alteration in the location of the biomes, large units of vegetation (e.g.: savannas, tropical forests, tundras, etc.).

Biome triangle classified by latitude, altitude and humidity (Author: Peter Halasaz).

On the other hand, there is an upward trend in the distribution of species in the high latitudes and a detriment in the lower latitudes. This has serious associated problems; the change in the species distribution affects their conservation and genetic diversity. Consequently, the marginal populations in lower latitudes, which have been considered very important for the long-term conservation of genetic diversity and due their evolutionary potential, are threatened by this diversity loss. And conversely, the populations in high latitudes would be affected by the arrival of other competing species that could displace those already present, being as invasive.

Species distribution

Within the scenario of climate change, species have some ability to adjust their distribution and to adapt to this.

But, what type of species may be responding more quickly to this change? It appears that those with a faster life cycle and a higher dispersion capacity will be showing more adaptability and a better response. This could lead to a loss of some plants with slower rates.

The Purple milk Thistle (Galactites tomentosa) is a plant with a fast life cycle and high distribution capacity  (Author: Ghislain118).

One factor that facilitates adjustment in the distribution is the presence of wildlife corridors: these are parts of the geographical area that enable connectivity and movement of species from one population to another. They are important because they prevent that some species can remain isolated and because they can also allow the movement to new regions.

Another factor is the altitudinal gradient, which provides shelter for many species, facilitates the presence of wildlife corridors and permits redistribution of species along altitude. Therefore, in those territories where there is greater altitudinal range, the conservation is favored.

In short, the ability of species to cope with climate change depends on the plant characteristics and the territory attributes. And, conversely, the species vulnerability to climate change occurs when the speed to displace their distribution or adapt their lives is less than the climate change velocity.

At internal level

Climate change also affects the plant as an organism, as it causes changes in their metabolism and phenology (periodic or seasonal rhythms of the plant).

One of the effects that pushes the climate change is the carbon dioxide (CO₂) concentration increase in the atmosphere. This could produce a fertilization phenomenon of vegetation. Due the CO₂ increase in the atmosphere it also increases the uptake by plants, thus increasing the photosynthesis and allowing greater assimilation. But, this is not all advantages, because for this an important water loss occurs due that the stomata (structures that allow gas exchange and transpiration) remain open long time to incorporate CO₂. So, there are opposing effects and fertilization will depend on the plant itself, but the local climate will also determine this process. Many studies have shown that various plants react differently to the CO₂ increase, since the compound affects various physiological processes and therefore there are not unique responses. Then, we find a factor that alters the plant metabolism and we cannot predict what will be the effects. Furthermore, this fertilizer effect is limited by the nutrients amount and without them production slows.

Photosynthesis process (Author: At09kg).

On the other hand, we must not forget that climate change also alters the weather and that this affects the vegetation growth and its phenology. This can have even an impact on a global scale; for example, could produce an imbalance in the production of cultivated plants for food.

PLANTS AS CLIMATE REGULATORS

Although one cannot speak of plants as regulators of global climate, it is clear that there is a relationship between climate and vegetation. However, this relationship is somewhat complicated because the vegetation has both effects of cooling and heating the weather.

The vegetation decreases the albedo; dark colours absorb more solar radiation and, in consequence, less sunlight is reflected outward. And besides, as the plants surface is usually rough, the absorption is increased. Consequently, if there is more vegetation, local temperature (transmitted heat) intensifies.

But, on the other hand, by increasing vegetation there is more evapotranspiration (set of water evaporation from a surface and transpiration through the plant). So, the heat is spent on passing the liquid water to gas, leading to a cooling effect. In addition, evapotranspiration also helps increase local rainfall.

Biophysical effects of different land uses and its consequences on the local climate. (From Jackson et al. 2008. Environmental Research Letters.3: article 0440066).

Therefore, it is an ambiguous process and in certain environments the cooling effect outweighs, while in others the heating effect has more relevance.

MITIGATION

Nowadays, there are several proposals to reduce climate change, but, in which way can the plants cooperate?

Plant communities can act as a sinks, carbon reservoirs, because through CO₂ assimilation, they help to offset carbon emissions. Proper management of agricultural and forest ecosystems can stimulate capture and storage of carbon. On the other hand, if deforestation were reduced and protection of natural habitats and forests increased, emissions would be diminished and this would stimulate the sink effect. Still, there is a risk that these carbon sinks may become emission sources; for example, due to fire.

Finally, we must introduce biofuels: these, unlike fossil fuels (e.g. petroleum), are renewable resources, since they are cultivated plants for use as fuels. Although they fail to remove CO₂ from the atmosphere or reduce carbon emissions, they get to avoid this increase in the atmosphere. For this reason, they may not become a strict mitigation measure, but they can keep neutral balance of uptake and release. The problem is that they can lead to side effects on social and environmental level, such as increased prices for other crops or stimulate deforestation to establish these biofuel crops, what should not happen.

Sugarcane crop (Saccharum officinarum) in Brazil to produce biofuel (Author: Mariordo).

REFERENCES

• Notes of Plant Physiology, Science of the Biosphere and Analysis of vegetation, Degree of Environmental Biology, UAB
• Hample & R. J. Petit. 2005. Conserving biodiversity under climate change: the rear edge matters. Ecology letters 8 (5): 461-467.
• Botanic Gardens Conservation International. 2015. How does climate change affect plants?. https://www.bgci.org/climate/climate_change_effects/
• Earth Observatory. 2015. How plants can change our climate. http://earthobservatory.nasa.gov/Features/LAI/LAI2.php

 

Evolution for beginners

Biological evolution is still not well understood by general public, and when we speak of it in our language abound expressions that confuse even more how mechanisms that lead to species diversity work. Through questions you may have ever asked yourself, in this article we will have a first look at the basic principles of evolution and debunk misconceptions about it.

IS EVOLUTION REAL? IT IS NOT JUST A THEORY OR AN IDEA WITHOUT EVIDENCES?

Outside the scientific field, the word “theory” is used to refer to events that have not been tested or assumptions. But a scientific theory is the explanation of a phenomenon supported by evidence resulting from the application of the scientific method.

The scientific method. Image by Margreet de Heer.

Theories can be modified, improved or revised if new data don’t continue to support the theory, but they are always based on some data, repeatable and verifiable experiments by any researcher to be considered valid.

So few people (sic) doubts about the heliocentric theory (the Earth rotates around the Sun), or the gravitational theory of Newton, but in the popular imagination some people believe that the theory of evolution made by Charles Darwin (and Alfred Russell Wallace) is simply a hypothesis and has no evidence to support it. With new scientific advances, his theory has been improved and detailed, but more than 150 years later, nobody has been able to prove it wrong, just the contrary.

WHAT EVIDENCE WE HAVE THAT EVOLUTION IS TRUE?

We have many evidences and in this post we will not delve into them. Some of the evidence available to us are:

• Paleontological record: the study of fossils tell us about the similarities and differences of existing species with others thousands or millions old, and to establish relationships respect each other.
• Comparative anatomy: comparison of certain structures that are very similar between different organisms, can establish whether they have a common ancestor (homologous structures, for example, five fingers in some vertebrates) if they have developed similar adaptations (analogous structures, for example, the wings of birds and insects), or if they have lost their function (vestigial organs, such as the appendix).

Homologous organs in humans, cats, whales and bats

• Embryology: the study of embryos of related groups shows a strong resemblance in the earliest stages of development.
• Biogeography: The study of the geographical distribution of living beings reveals that species generally inhabit the same regions as their ancestors, although there are other regions with similar climates.
• Biochemistry and genetics: chemical similarities and differences allow to establish relationships among different species. For example, species closely related to each other have a structure of their DNA more similar than others more distant. All living beings share a portion of DNA that is part of your “instructions”, so there are also found in a fly, a plant or a bacterium, proof that all living things have a common ancestor.

IS IT TRUE THAT ORGANISMS ADAPT TO THE ENVIRONMENT AND ARE DESIGNED FOR LIVING IN THEIR HABITAT?

Both expressions, frequently used, mean that living beings have an active role to adapt to the environment or “someone” has designed them to live exactly where they are. It is a typical example of Lamarck and giraffes: as a result of stretching the neck to reach the higher leaves of the trees,  currently giraffes have this neck for giving it this use. They have a necessity, they change their bodies to success. It is precisely upside down: it is the habitat that selects the fittest, nature “selects” those that are most effective to survive, and therefore reproduce. It is what is known as natural selection, one of the main mechanisms of evolution. It needs three requirements to act:

• Phenotypic variability: there must be differences between individuals. Some giraffes necks were slightly longer than others, just as there are taller people than others, with blue or brown eyes.
• Biological fitness: this difference has to suppose an advantage. For example, giraffes with a slightly longer neck could survive and reproduce, while the others don’t.
  
• Heredity: these characters must be transmitted to the next generation, the offspring will be slightly different to that feature, while “short neck” feature transmits less and less.

The variability in the population causes individuals with favorable characteristics to reproduce more and pass on their genes to the next generation, increasing the proportion of those genes. Image taken from Understanding evolution

Over the years these changes are accumulated until the genetic differences are so big that some populations may not mate with others: a new species has appeared.

If you thought that this is similar to artificial selection that we do with the different breeds of dogs, cows who give more milk, trees bearing more fruit and larger, congratulations, you think like Darwin as it was inspired by some of these facts. Therefore, living beings are mere spectators of the evolutionary process, depending of changes in their habitat and their genetic material.

WHY ORGANISMS ARE SO DIVERSE?

Genetic variability allows natural selection act. Changes in the genetic material (usually DNA) are caused by:

• Mutations: changes in the genome that may be adverse or lethal for survival, indifferent or beneficial to survival and reproduction. If they have benefits, they will pass to the next generations.
• Gene flow: is the motion of genes between populations (migration of individuals allows this exchange when mate with others in a different population).
• Sexual reproduction: allows recombination of genetic material of different individuals, giving rise to new combinations of DNA.

Populations that have more genetic variability are more likely to survive if happen any changes in their habitat. Populations with less variability (eg, being geographically isolated) are more sensitive to any changes in their habitat, which may cause their extinction.

Evolution can be observed in beings with a very high reproduction rate, for example bacteria, since mutations accumulate more quickly. Have you ever heard that bacteria become resistant to our antibiotics or some insects to pesticides? They evolve so quickly that within a few years were selected the fittest to survive our antibiotics.

ARE WE THE MOST EVOLVED ANIMALS?

Theory of Evolution has various consequences, such as the existence of a common ancestor and that therefore, that we are animals. Even today, and even among the young ones, there is the idea that we are something different between living beings and we are in a special podium in the collective imagination. This anthropocentric thinking caused Darwin mockery and confrontations over 150 years ago.

Caricature of Darwin as an orangutan. Public domain image first published in 1871

We use our language to be “more evolved” as a synonym for more complex, and we consider ourselves one species that has reached a high level of understanding of their environment, so many people believe that evolution has come to an end with us.

The question has a mistake of formulation: actually evolving pursues no end, it just happens, and the fact that millions of years allows the emergence of complex structures, it does not mean that simpler lifeforms are not perfectly matched in the habitat where they are. Bacteria, algae, sharks, crocodiles, etc., have remained very similar over millions of years. Evolution is a process that started acting when life first appeared and continues to act in all organisms, including us, although we have changed the way in which natural selection works  (medical and technological breakthroughs, etc.).

SO IF WE COME FROM MONKEYS, WHY DO STILL MONKEYS EXIST?

The truth is that we don’t come from monkeys, we are monkeys, or to be more rigorous, apes. We have not evolved from any existing primate. As we saw in a previous post, humans and other primates share a common ancestor and natural selection has been acting differently in each of us. That is, evolution has to be viewed as a tree, and not as a straight line, where each branch would be a species .

First scheme of the evolutionary tree of Darwin in his notebook (1837). Public domain image.

Some branches stop growing (species become extinct), while others continue to diversify. The same applies to other species, in case you have asked yourself, “if amphibians come from fish, why are there still fish?”. Currently, genetic analyzes have contributed so much data that they make so difficult to redesign the classical Dariwn’s tree.

Classification of live organisms based on the three domains Archaea, Bacteria and Eukarya, data of Carl R. Woese (1990). Included in Eukarya there are the Protista, Fungi, Plantae and Animalia kingdoms. Image by Rita Daniela Fernández.

Evolution is a very broad topic that still generates doubts and controversies. In this article we have tried to bring to uninitiated people some basics, where we can delve into the future. Do you have any questions about evolution? Are you interested into a subject that we have not talked about? You can leave your comments below.

REFERENCES

• Darwin’s Tree of Life is a Tangled Bramble Bush
• Algunas reflexiones sobre la clasificación de los seres vivos
• Museos vivos. Evidencias de la evolución
• Las ideas en la ciencia: Teoría, hipótesis y leyes
• Frequently asked questions about evolution
• Brock et. al. 1999. Biología de los microorganismos. Ed. Prentice Hall.
• Massa, Renato. El origen de la vida. La evolución de las especies. Ed. Susaeta.
• Cover image