Tuesday, December 26, 2017

Rhizome

Rhizome Definition


A rhizome (also known as rootstocks) is a type of plant stem situated either at the soil surface or underground that contains nodes from which roots and shoots originate (shown below). Rhizomes are unique in that they grow perpendicular, permitting new shoots to grow up out of the ground. When separated, each piece of a rhizome is capable of producing a new plant.


Zingiber officinale fresh rhizome


Rhizome Function


The primary function of the rhizome is the storage of nutrients, including carbohydrates and proteins, until the plant requires them for the growth of new shoots or to survive the winter in a process termed vegetative reproduction. Farmers use vegetative reproduction to laterally propagate plants such as hops, ginger and various grass species. Some rhizomes are also consumed or used as a seasoning, including ginger and turmeric.


Rhizome Examples


Underground Rhizomes


By far the most dominant type of rhizome is the underground rhizome (pictured below), which is situated underground and includes ginger, hops, poison oak, grass species, and bamboo. Many of these plants have rhizomes that are consumed by humans (e.g., ginger).


Jiaogulan Rhizome


Above-ground Rhizomes


While most rhizomes are situated underground, some plants have rhizomes that grow at the soil level or above (shown below). Examples of these plant species include ferns and irises.


Golden Chicken Fern


Multi-layered Rhizomes


The majority of rhizomes occur as a single layer from which shoots and roots originate. However, there are some plant species which form multiple layers in a complex network (e.g., Giant Horsetails [shown below]).


the Giant Horsetail


Quiz


1. True or False, rhizomes are always found underground.

Answer to Question #1

2. Which of the following statements is TRUE regarding the function of the rhizome:
A. The rhizome is an enzyme found in plants
B. The function of the rhizome is to store nutrients
C. The function of the rhizome is to provide a defense against pathogens
D. None of the above are true

Answer to Question #2

References



  • Holshouser D, Chandler J, and Wu H. (1996). Temperature-Dependent Model for Non-Dormant Seed Germination and Rhizome Bud Break of Johnsongrass (Sorghum halepense). Weed Science. 44(2): 257-265.

  • Kraemer G and Alberte R. (1993). Age-related patterns of metabolism and biomass in subterranean tissues of Zostera marina (eelgrass). Marine Ecology Progress Series. 95: 193-203.

  • Marba N and Duarte C. (1998). Rhizome elongation and seagrass clonal growth. Marine Ecology Progress Series. 174: 269-280.

  • McSteen P. (2009). Hormonal Regulation of Branching in Grasses. Plant Physiology. 149 149 (1) 46-55. DOI: 10.1104/pp.108.129056

  • Shaver G and Billings W. (1976). Carbohydrate Accumulation in Tundra Graminoid Plants as a Function of Season and Tissue Age. Flora. 165(3): 247 – 267.



Rhizome

Keystone Species

Keystone Species Definition


Keystone species are those which have an extremely high impact on a particular ecosystem relative to its population. Keystone species are also critical for the overall structure and function of an ecosystem, and influence which other types of plants and animals make up that ecosystem. Thus, in the absence of a keystone species, many ecosystems would fail to exist. A common example of keystone species in the context of conservation biology is the predator-prey relationship. Small predators that consume herbivorous species prevent such herbivores from decimating the plant species in the ecosystem, and are considered keystone species. In this scenario, despite the low number of predators required to maintain a low population of herbivorous species, without this keystone species, the herbivore population would continue to grow, and thus consume all of the dominant plant species in the ecosystem.


Keystone Species Examples


Sea Otter


The sea otter (shown below) is considered a keystone species as their consumption of sea urchins, preventing the destruction of kelp forests caused by the sea urchin population. Kelp forests are a critical habitat for many species in nearshore ecosystems. In the absence of sea otters, sea urchins feed on the nearshore kelp forests, thereby disrupting these nearshore ecosystems. However, when sea otters are present, their consumption of sea urchins restricts the sea urchin population to smaller organisms confined to protective crevices. Thus, the sea otter protects the kelp forests by reducing the local sea urchin population.


Sea Otter


Large Mammalian Predators


While small predators are important keystone species in many ecosystems, as mentioned above, large mammalian predators are also considered keystone species in larger ecosystems. For example, the lion, jaguar (shown below), and gray wolf are considered keystone species as they help balance large ecosystems (e.g., Central and South American rainforests) by consuming a wide variety of prey species.


Black jaguar


Sea Star


Sea stars (shown below) are another commonly recognized keystone species as they consume mussels in areas without natural predators. In many cases, when the sea star is removed from an ecosystem, the population of mussels proliferates uncontrollably, and negatively effects the resources available to other species within the ecosystem.


Sea Star


Quiz


1. Which of the following statements is TRUE regarding keystone species?
A. Keystone species are usually herbivores.
B. Keystone species are usually predators.
C. Keystone species are the most abundant species in an ecosystem.
D. Keystone species are non-essential species in an ecosystem.

Answer to Question #1

2. Which of the following statements is FALSE regarding keystone species?
A. Without a keystone species, many ecosystems would fail to exist.
B. Keystone species populations are typically small compared to other species in an ecosystem.
C. Kelp forests are considered a keystone species.
D. Sea otters, lions, and sea stars are also considered important keystone species.

Answer to Question #2

References



  • Bucci et al. (2017). Sea Star Wasting Disease in Asterias forbesi along the Atlantic Coast of North America. PLoS One. 12(12):e0188523.

  • Gooding, R and Harley, C. (2015). Quantifying the Effects of Predator and Prey Body Size on Sea Star Feeding Behaviors. Biol Bull. Jun;228(3):192-200.

  • Humphries et al. (2017). To Everything There Is a Season: Summer-to-Winter Food Webs and the Functional Traits of Keystone Species. Integr Comp Biol. 57(5):961-976. doi: 10.1093/icb/icx119.

  • Petes eta l. (2008). Effects of environmental stress on intertidal mussels and their sea star predators. Oecologia 156(3):671-80. doi: 10.1007/s00442-008-1018-x.



Keystone Species

Coevolution

Coevolution Definition


In the context of evolutionary biology, coevolution refers to the evolution of at least two species, which occurs in a mutually dependent manner. Coevolution was first described in the context of insects and flowering plants, and has since been applied to major evolutionary events, including sexual reproduction, infectious disease, and ecological communities. Coevolution functions by reciprocal selective pressures on two or more species, analogous to an arms race in an attempt to outcompete each other. Classic examples include predator-prey, host-parasite, and other competitive relationships between species. While the process of coevolution generally only involves two species, multiple species can be involved. Moreover, coevolution also results in adaptations for mutual benefit. An example is the coevolution of flowering plants and associated pollinators (e.g., bees, birds, and other insect species).


Coevolution Examples


Predator-Prey Coevolution


The predator-prey relationship is one of the most common examples of coevolution. In this respect, there is a selective pressure on the prey to avoid capture and thus, the predator must evolve to become more effective hunters. In this manner, predator-prey coevolution is analogous to an evolutionary arms race and the development of specific adaptations, especially in prey species, to avoid or discourage predation.


Herbivores and plants


Similar to the predator-prey relationship, another common example of coevolution is the relationship between herbivore species and the plants that they consume. One example is that of the lodgepole pine seeds, which both red squirrels and crossbills eat in various regions of the Rocky Mountains. Both herbivores have different tactics for extracting the seeds from the lodgepole pine cone; the squirrels will simply gnaw through the pine cone, whereas the crossbills have specialized mandibles for extracting the seeds. Thus, in regions where red squirrels are more prevalent, the lodgepole pine cones are denser, contain fewer seeds, and have thinner scales to prevent the squirrels from obtaining the seeds. However, in regions where crossbills are more prevalent, the cones are lighter and contain thick scales, so as to prevent the crossbills from accessing the seeds. Thus, the lodgepole pine is concurrently coevolving with both of these herbivore species.


Acacia ants and Acacias


An example of coevolution that is not characteristic of an arms race, but one which provides a mutual benefit to both a plant species and insect is that of the acacia ants and acacia plants. In this relationship, the plant and ants have coevolved to have a symbiotic relationship in which the ants provide the plant with protection against other potentially damaging insects, as well as other plants which may compete for nutrients and sunlight. In return, the plant provides the ants with shelter and essential nutrients for the ants and their growing larvae (shown below).


Ant - Pseudomyrmex species, on Bull Thorn Acacia


Flowering Plants and Pollinators


Another example of beneficial coevolution is the relationship between flowering plants and the respective insect and bird species that pollinate them. In this respect, flowering plants and pollinators have developed co-adaptations that allow flowers to attract pollinators, and insects and birds have developed specialized adaptations for extracting nectar and pollen from the plants (shown below).


Prima Vera


Research indicates that there are at least three traits that flowering plants have evolved to attract pollinators:



  • Distinct visual cues: flowering plants have evolved bright colors, stripes, patterns, and colors specific to the pollinator. For example, flowering plants seeking to attract insect pollinators are typically blue an ultraviolet, whereas red and orange are designed to attract birds.

  • Scent: flowering plants use scents as a means of instructing insects as to their location. Since scents become stronger closer to the plant, the insect is able to hone-in and land on that plant to extract its nectar.

  • Some flowers use chemical and tactile means to mimic female insect species to attract the male species. For example, orchids secrete a chemical that is the same as the pheromones of bee and wasp species. When the male insect lands on the flower and attempts to copulate, the pollen is transferred to him.


Hummingbirds are another type of pollinator that have coevolved for mutual benefit. The hummingbirds serve as pollinators and the flowers supply the birds with nutrient-rich nectar. The flowering plants attract the hummingbirds with certain colors, the shape of the flower accommodates the bird’s bill, and such flowers tend to bloom when hummingbirds are breeding. Coevolution of such flowering plants with various hummingbird species is evident by the distinct shape and length of the flower’s corolla tubes, which have adapted to the shape and length of the hummingbird bill that pollenates that plant. The shape of the flower has also adapted such that the pollen becomes attached to a particular region of the bird while it consumes the nectar from the flower (shown below).


Hummingbird seen on the Sonoran Desert


Quiz


1. Which of the following statements is TRUE regarding coevolution?
A. Coevolution can result in a symbiotic or mutually beneficial relationship between two species.
B. Coevolution can be the result of selective pressures between two species, resulting in an arms race between them.
C. Both A and B are correct
D. None of the above

Answer to Question #1

2. Which of the following is NOT an example of coevolution?
A. Acacia ants and lodgepole pines
B. Acacia ants and acacia plants
C. Crossbills and lodgepole pines
D. Red squirrels and lodgepole pines

Answer to Question #2

3. An example of coevolution for mutual benefit is:
A. Red Squirrels and lodgepole pines
B. Crossbills and lodgepole pines
C. Large mammalian predators (e.g., foxes, wolves) and hedgehogs or skunks
D. Acacia ants and acacia plants

Answer to Question #3

References



  • Benkman et al. (2003). Reciprocal Selection Causes a Coevolutionary Arms Race between Crossbills and Lodgepole Pine. The American Naturalist. June.

  • Endara et al. (2017). Coevolutionary arms race versus host defense chase in a tropical herbivore– plant system. PNAS. 114(36): E7499-E7505.

  • Eriksson, Ove. (2016). Evolution of angiosperm seed disperser mutualisms: the timing of origins and their consequences for coevolutionary interactions between angiosperms and frugivores. Biological Reviews. 91(1): 168-186.

  • Pauw, A. et al. (2017). Long-legged bees make adaptive leaps: linking adaptation to coevolution in a plant–pollinator network. Proceedings of the Royal Society B. September. DOI: 10.1098/rspb.2017.1707

  • Janzen, D. (1966). Coevolution of mutualism between ants and acacias in Central America. Evolution. 20(3): 249-275.

  • Smith, J and Benkman, C. (2006). A Coevolutionary Arms Race Causes Ecological Speciation in Crossbills. The American Naturalist. November.



Coevolution