Angiosperms Part Two


 Lifecycle & Ecology


https://www.youtube.com/watch?v=j6LCCxFeDq8&feature=youtube_gdata

The Angiosperm Life Cycle Video – This is a look at the growth, fertilization, and pollination events in flowering plants. The diversity of morphology in angiosperm structures helps to facilitate this life cycle. It is the reason that angiosperms have been able to be ubiquitous throughout the various biomes of our Earth! Created by Emily Thomas.

Angiosperms are considered to be one of the greatest examples of symbionts in nature, due to their many mutualistic relationships with pollinators, fungi, herbivores and others. They can be found in almost any environment, so long as there is sunlight, some form of water, and a way to spread their offspring. The general life cycle of angiosperms is explained in the video on this page, ‘Angiosperm Life Cycle.’ While this video focuses on a more general view, the four major events that make up flowering plant reproduction (pollen development, egg development, pollination and fertilization) will be focused on in more detail here.  Angiosperms produce two types of spores; microspores which lead to the generation of pollen and megaspores which form the structure that houses female gametophytes (Boundless, 2014).

Pollen develops inside the stamen. Inside the anther of an angiosperm lie the diploid microspores. These microspores undergo meiosis to become haploid microspores. Haploid microspores then undergo mitosis to develop into pollen grains. Pollen forms from the male gametophyte in flowering plants.  The gametophyte consists of two cell types: tube cells to aid in fertilization and generative cells which generate sperm cells (Taiz, 2006).

Egg development occurs inside the carpel. Inside of the ovaries are egg producing structures known as ovules. Inside of an ovule are diploid cells. Similar to the production of pollen, these diploid cells divide via meiosis to become haploid cells that are the megaspores. These spores then go through three rounds of mitosis forming seven cells. One of the cells has two nuclei, called the endosperm and another of the cells becomes the egg. The remaining five cells are not used in reproduction (Boundless, 2014).

Once both eggs and pollen development have taken place they are ready for pollination. Pollination can occur in many ways; two major forms are wind and water dispersal. Unique to angiosperms is the use of pollinators such as birds and bees.  The last step in flowering plant reproduction is fertilization. For fertilization to occur the tube cell of the male gametophyte creates a tube to the ovule (Taiz,  2006). The generative cell uses this tube to send sperm down to the ovule so fertilization can occur. One sperm will fuse with the egg forming a zygote the other fuses with the endosperm forming a triploid endosperm cell. This process, known as double fertilization, is unique to angiosperms (Derksen, 2013). The endosperm later develops into nutrient tissue while the zygote divides by mitosis, developing into an embryo which grows into a mature plant.

Angiosperms spend most of there life in the adult stage known as a sporophyte. When we see trees, grass, flowers, vegetables in a garden we are seeing sporophytes! Angiosperms are very important due to their abundance and impact on almost every habitat on earth. Due to their diverse morphology they can range from the small to massive, aquatic to mountainous, grass to trees and everything in between.

A fruiting angiosperm. Photo taken by Nick White.

A fruiting angiosperm. Photo by Nick White.


 Evolutionary History


 

The Amorphophallus titanum, or Corpse flower is one of the most bizarre and oldest ancestors of modern angiosperms. Photo taken by Nick White

The Amorphophallus titanum, or Corpse flower is one of the oldest and most bizarre ancestors of modern angiosperms. Photo by Nick White.

Not surprisingly, angiosperms are the most commonly found type of land plant. Angiosperms evolved in the Cretaceous era, around the same time as many groups of modern insects. Many of these insects acted as pollinators that drove the evolution of both angiosperms and the insects themselves. (Soltis 2005). Due to the availability of pollinators (insects occupy nearly every environment on the planet) it has allowed angiosperms to become the most numerous plant found on land. This relationship is considered one of the greatest examples of symbioses in nature due to their many mutualistic relationship with pollinators, fungi, herbivores and others. They can be found in almost any environment, so long as there is sunlight, some form of water, and a way to spread their offspring.With co-evolution, these two species have been able to occupy places that few other species previously could, changing the habitats of the entire planet.(Lerner 2008)

Unlike many land plants, angiosperms did not evolve from gymnosperms. It is unclear what type of plants gave rise to angiosperms. (Angiosperms 2014) Some scientists believe that a group of plants known as “seed ferns” ,or pteridosperms, may have been the progenitor of the angiosperms. These “seed ferns” were around for many millions of years before angiosperms and yet have similar traits like seed-bearing capsules and specialized organs that produced pollen. While we are still not exactly sure how ancient angiosperms may have come about, we have an idea of what these ancestors may have looked like. They were likely small with small flowers. The flowers were probably green and not at all like the flowers we are used to as their sepals and petals would not be separated or distinguishable (Angiosperms 2014). And while the exact way that angiosperms evolved to what we know today is still unclear, their impact on our world today is obvious.

 


 A Survey of Extant Diversity


Angiosperms are arguably the largest extant group of plants on the planet today.  At least 260,000 living species exist, which are classified into 453 families (Soltis 2005). The most popular

The photo above are barrel cacti (Echinocactus grusonii) displaying the numerous spines that are used for protection. Additionally, these cacti have modified leaves, which help retain water in dry environments; they are also known as succulents. Photo by Nick White.

lineage would be the eudicots, which includes most flowering plants.  Some other major lineages are the Monocotyledons, containing families like lilies, grasses, and orchids, and the Nymphaeaceae, which hold the water lilies and their relatives (Soltis 2005).  Angiosperms inhabit all seven continents, as well as the oceans.  They are able to occupy just about any environment on earth, for example, high mountaintops, deep oceans, freezing tundras, and of course, warm, wet rainforests.  Their abundance in these environments is immense.  They have an extremely large genome, which may explain their ability to exist in so many different morphological forms. Some examples of these forms include grasses, climbing vines, large trees, and small flowers.  By diversifying their physiology, angiosperms have been able to adapt to the variety of ecosystems which cover the earth. The cactus, for example, has modified leaves, called spines, which help to prevent it’s desiccation in dry, arid deserts (see photo below).  Some types of angiosperms can be quite special and complex in terms of their nutrient acquisition, for example, carnivorous flowers, or poisonous vines.

Angiosperms provide an enormous environmental and economical importance.  Environmentally, they use the carbon dioxide we produce, and turn it into the oxygen that is pertinent to our survival.  Obviously, they also provide food for a variety of organisms, including humans.  All of the fruits and vegetables bought in our grocery stores are products of angiosperms.  Many insects also feed on these plants leaves, and bees use them to create their honey.  Trees provide shelter and places to build homes for countless organisms, such as birds and squirrels, while we use the wood to build our houses,  and make our paper.  In fact, the clothes we wear everyday come from cotton plants, which are angiosperms.  Certain angiosperms are also used as a source to create medicines.  A common medicine, morphine, is made from the opium poppy (Papaver somniferum), and is used everyday in hospitals for pain relief (Taylor 1996).  Another drug called cynarin comes from a chemical in the common artichoke (Cynara scolymus).  It is being used in Germany to treat liver problems and hypertension (Taylor 1996).  As you might have guessed, the abundance of angiosperms is crucial for human existence, as well as the majority of other organisms on earth, and it would be impossible to name every use and importance of these plants.


 Conclusion


Researchers are working to clarify the emergence of angiosperms and delineate their origins to compensate for discrepancies between the fossil and molecular clock data (Peppe, 2013).  Like many fossil records, the angiosperm fossil record is believed to lag behind the time of divergence for the clade. Peppe explains that the only way to resolve the issues between the fossil record, which suggests the arrival of angiosperms in the early Cretaceous period, and molecular dating, which suggests arrival in the Jurassic,  is to look for Triassic and Jurassic fossils “with an eye toward finding angiosperm and angiosperm-like plant fossils” (2013).

One such study, the Hochuli and Feist-Burkardt (2013), examined fossilized pollen samples to try to identify early angiosperms and potential features which can be used to firmly identify the clade. The research identified the pollen as “Triassic and Jurassic angiosperm-like fossils,” which, while not angiosperms themselves, could be useful in establishing ancestral features and pinpointing groups which were evolving traits useful in characterizing modern angiosperms. By finding these earlier emerging pre-angiosperm groups within the fossil records, scientists can develop better hypotheses about when and where the earliest angiosperm fossils may be found (Peppe, 2013).

Other areas of ongoing research are expansive. The worldwide prominence of angiosperms has led to curiosity surrounding their reproduction, diversity, speciation, and uses. The diversity and accessibility of angiosperms means that funding availability tends to be the determining factor in driving research. As a result, much research focuses on the medicinal and agricultural uses of the flowering plants, because of the implications for humans (Reddy and Yang, 2011). Angiosperms include everything from corn to oak trees, so research focuses on effective crop cultivation, pesticide use, sustainability, and industrial uses. One of the most interesting research topics in agriculture surrounds the introduction of genetically modified organisms (GMOs) to the market (Miraglia, et. al, 2004).

From an ecological perspective, angiosperms reproduction via pollination and their intrinsic link to their pollinators has driven many research projects on the coevolution of plants and animals. The wide range of shape, size, color, and chemical secretions of the plants’ flowering portions, as well as the fast morphological differences in their fruiting body, have led to morphological specificity and behavioral patterning amongst insects, birds, and some mammals (Jarzden and Dilcher, 2010).


 Additional Resources


The angiosperm group is a diverse one. Flowering plants, which make up so much of what we see, eat, and use every day, are a source of fascination. These plants have adapted to inhabit nearly every corner of land on the Earth. Curiosity surrounding the variation, morphology, evolution, and prevalence of angiosperms, has led to the establishment of many resources for those looking to further their understanding. The following are just some of the many videos and articles available for continued learning about these magnificent plants.

Angiosperm In Encyclopædia Britannica (http://www.britannica.com/EBchecked/topic/24667/angiosperm) is a very comprehensive encyclopedia entry that covers everything from general features, to reproduction, to classification, and fossilization. The entry is broken into subsections so that readers can focus on their areas of interest or questions. There are even quizzes to check understanding,

Biology for Kids (http://www.biology4kids.com/files/plants_angiosperm.html) provides a shorter, basic introduction to angiosperms as a whole in a language that is accessible to explorers of all ages. The page also provides resources for further learning.

‘Sexual Reproduction in Flowering Plants’ and ‘Flowers: Sexual Reproduction in Flowering Plants,’ which can be found on YouTube at https://www.youtube.com/watch?v=w1BSCJrH4lU and https://www.youtube.com/watch?v=hf9XlqXcal0 , help to explain the reproductive parts of a flower and the mechanisms surrounding pollination, fertilization, and fruit development. Both videos are about 2 and a half minutes long and appropriate for an audience with a basic understanding of plants.

The Pollinator Partnership website (http://www.pollinator.org/) is a great resource for those curious about learning more about the organisms which serve as pollen carriers for angiosperms. It not only provides basic information and further resources, but also explains the ecological importance of pollinators and how we can protect the creatures which help us sustain our agriculture and industry.

Flowering Plants: Keys to Earth’s Evolution and Human Well-Being is a 2005 Q&A interview with Pamela Soltis Ph.D, a key contributor to the Tree of Life. Soltis describes, in an easy and engaging way, the value of angiosperms in terms of their diversity, uses, and everyday influence on humans. It answers a lot of the “why should we care?” questions and explains the intertwined relationship of animal and plant. The full transcript can be found at http://www.actionbioscience.org/genomics/soltis.html.


Works Cited


Angiosperms. (2008). In L. Lerner & B. Lerner (Eds.), The Gale Encyclopedia of Science (4th ed., Vol. 1, p. 217). Detroit: Gale.Carter, J. (2014, January 17). Angiosperms. Retrieved March 6, 2015, from http://biology.clc.uc.edu/courses/bio106/angio.htm

Boundless. “Evolution of Angiosperms.” Boundless Biology. Boundless, 14 Nov. 2014. Retrieved 29 Mar. 2015 from https://www.boundless.com/biology/textbooks/bound20-11841/

Carter, J. Stein. (2014, Jan. 17) Angiosperms. Retrieved from http://biology.clc.uc.edu/courses/bio106/angio.htm.

Derksen, J., & Pierson, E. (2013, September 10). Life cycles. Retrieved March 30, 2015 from http://www.vcbio.science.ru.nl/en/virtuallessons/pollenreproduction/.

Dilcher, D. (2000). Toward a new synthesis: Major evolutionary trends in the angiosperm fossil record. Proceedings of the National Academy of Sciences, 7030-7036. http://dx.doi.org/10.1073/pnas.97.13.7030

Hedges, S., & Kumar, S. (2009). Plants. In The Timetree of Life (pp. 133-137, 162-165). Oxford: Oxford University Press.

Jarzen, David M. and Dilcher, David L. (2010). Coevolution between flowering plants and insect pollinators. In AccessScience. McGraw-Hill Education. Retrieved from http://accessscience.com/content/coevolution-between-flowering-plants-and-insect-pollinators/YB100138

Miraglia, M., Berdal, K. G., Brera, C., Corbisier, P., Holst-Jensen, A., Kok, E. J., & Zagon, J. (2004). Detection and traceability of genetically modified organisms in the food production chain. Food and Chemical Toxicology, 42(7), 1157-1180. http://dx.doi.org/10.1016/j.fct.2004.02.018

Peppe, D. (2013, October 15). What do we know about the origin of flowering plants? Retrieved March 30, 2015, from http://blogs.egu.eu/network/palaeoblog/2013/10/15/what-do-we-know-about-the-origin-of-flowering-plants/

Reddy, N., & Yang, Y. (2011). Potential of plant proteins for medical applications. Trends in biotechnology, 29(10), 490-498. http://dx.doi.org/10.1016/j.tibtech.2011.05.003

Soltis, D., Soltis, P., & Edwards, C. (2005, June 3). Angiosperms: Flowering Plants. Retrieved March 6, 2015, from http://tolweb.org/Angiosperms/20646/2005.06.03

Taylor, Leslie. (1996) Plant-Based Drugs and Medicines. Retrieved March 29, 2015, from http://www.rain-tree.com/plantdrugs.htm#.VRh1E4tAxgs.

Taiz, L., & Zeiger, E. (2006) Topic 1.3. Retrieved March 30, 2015, from           http://5e.plantphys.net/article.php?id=474

Angiosperms

From flower shops to the produce section at the supermarket angiosperms, and their by-products, can be seen everywhere. Comprised of more than 260,00 species the angiosperm taxon is extremely diverse. The most abundant of the green plant division, many of the most economically and agriculturally important plants are angiosperms. Their diversity has allowed them to colonize multiple different types of habits and survive in various environments across the world. Clovers, Sunflowers, and Zebra Succulent are three exemplary species for angiosperm diversity. Though they are diverse they share several features such as their unique reproduction morphology, which will be discussed in this blog.


Phylogenetic Tree of Life


Phylogeny of Angiosperms and its groups

Phylogeny of Angiosperms and it’s groups. Created by Alyssa Riddle.

There are four supergroups of Eukaryotes and they include the Unikonts, the Chromalveolates, the Excavates, and the Archeaplastida. Archeaplastida are also called Plantae, and is the supergroup that the angiosperms belong to.

Archeaplastida contains three major lineages including Glaucophytes (microalgae), Rhodophyta (red algae), and the lineage that contains angiosperms, the Green Plants (Hedges & Kumar, 2009). The lineage of land plants stem from the Green Plants and are known as the Embryophytes. Sixteen different lineages stem from the Embryophytes, but the group that the angiosperms belong to are the Spermatopsida. Spermatopsida contain groups such as the conifers, seed plants, and flowering plants (Hedges & Kumar, 2009).

Analysis in the last five years has led scientists to agree that Amborella is the base of the angiosperm’s evolutionary tree. Major groups that branch off from Amborella trichopoda are Nymphaeaceae (water lilies and relatives), Austrobaileyales, Magnoliids, Chloranthaceae, Ceratophyllaceae, Monocotyledons (lilies, orchids, grasses), and eudicots (most flowering plants).

The order of taxonomic hierarchy for angiosperms is ranked: Eukaryote, Archeaplastida, Green Plants, Embryophytes, Spermatopsida, Angiosperms. Angiosperms contain at least 260,000 living species which are classified into 453 families and over 904,649 species (Hedges & Kumar, 2009).

See photo gallery below for some examples of these species.

Above is a Photo Gallery exampling some species in order to show the wide range of diversity in Archaeplastida. (Photos by Alyssa Riddle)


 Fossil Evidence and the Molecular Clock 


Angiosperms are a specific group within the Plantae Kingdom.

This timeline represents the estimated divergence of the kingdom Plantae. This diagram displays the diversification of various lineages and their relationships to the Angiosperm clade. The timeline is based upon molecular clock data provided by Hedges, Blair, and Kumar through the Timetree of Life project (2009). Created by Emily Thomas.

Fossil and molecular clock evidence agree that angiosperms are the most recently evolved of the major groups of plants. Both bodies of evidence also agree that the clade diverged from their sister group the gymnosperms, the cone-bearing plants (“Angiosperms,” 2008).

The timing of this divergence is not fully resolved by the fossil record and molecular clock estimates. The lack of a comprehensive fossil record has led to molecular clock evidence as more widely accepted by the scientific community. This evidence suggests that angiosperms arose approximately 175 million years ago (Hedges & Kumar, 2009). The hypothesized phylogenetic and chronological relationships of angiosperms to gymnosperms, as well as the other plant lineages, based on molecular clock evidence, are see in the figure to the right. 

Angiosperm Fossil Evidence

The most definite evidence of angiosperms in the fossil record comes from Cretaceous era fossils are the most definite evidence . The fossil record of angiosperms display a wide variety of structures, shape, and size. The vast morphological diversity has made it difficult to resolve relationships between the major angiosperm clades, but shows early diversification of lineages (Soltis, Soltis, & Edwards, 2005)Fossilization of leaves, pollen, wood, and floral structures have allowed for character based analysis of evolution (Dilcher, 2000). While fossil evidence has provided a basic understanding of angiosperm diversity throughout time, scientists must rely on the combination of preserved specimen’s physical and genetic characteristics to develop a more definite understanding of the angiosperm clade and relationships among it’s lineages.

This timeline represents the estimated time of diversification of the angiosperm clade. Based on molecular clock data (Hedges & Kumar, 2009), the diagram shows the rapid diversification of angiosperms. This diversification occurred in a relatively short geological time frame (approx.. 40 million years). Created by Emily Thomas.

Molecular Clock

While molecular clock evidence is the most widely used for examining phylogenetic relationships, complications arise in using molecular clock evidence for plants because of inconsistent evolution rates among different lineages (Dilcher, 2000).

Molecular clock evidence predates fossilization records for angiosperms by approximately 50 million years (Soltis, et. al, 2005). This unifies the angiosperm clade as a monophyletic group, defined by one evolutionary event, but does not fully resolve relations between other plant lineages. (Hedges & Kumar, 2009). 

 Within the angiosperm clade there are 5 major extant groups (Eudicots, Ceratophyllales, Monocots, Magnoliid, Chloroanthales) and 3 other “primitive” (non-extant) groups (Austrobaileyales, Nymphaelales, and Amborellales) (Hedges & Kumar, 2009).

 The major divergences amongst these groups are represented in the phylogenetic timeline above. Molecular evidence suggests the first divergence within the clade was the Amborellales approximately 174.9 mya. The Nymphaeales diverged  approximately 167.3 mya. The Austrobaileyales  diverged 159.5 mya, the Chloroanthales 150.1 mya, and the Magnoliids 147.8 mya. The most recent divergences were of Monocots  146.6 mya, and the Ceratophyllales 146.3 mya (Hedges & Kumar, 2009). 

 


 Evolutionary Innovations


Over time, specific evolutionary features, have distinguished angiosperm reproduction. The development of non-exposed seeds, housed within a flower structure, defines the group. This evolutionary feature has led to an abundance of morphological variation and widespread distribution of this group. Angiosperm flower structures have evolved in response to ecological pressures rapidly, and this success has led to the group’s survival, nearly universally, across the diverse ecosystems of our planet (Carter 1997).

 Angiosperms produce their gametes in separate organs from their bodies and these are generally housed in a flower. Fertilization takes place in structures to keep the process relatively unexposed to the elements. Flowering plants are the most diverse organism on the planet after insects.

Spider Wasp, under a dissection microscope. This organism is a common pollinator and of the family Pompilidae. Photo by Nick White.

Flowers come in an astounding number of colors, shapes, sizes, arrangements, and smells. All of these are evolutionary innovations which assist in attracting pollinators. Attraction is effected by color, scent, and the production of nectar, which may be secreted in some part of the flower. Pollinator’s relationship with their flowers are a textbook example of coevolution, as some animals evolve specifically to cater to a flowers pollination needs. These animals transport the flowers pollen to a wider geographic range to give them an excellent diversity within the population. (Carter, 1997)

Flower organs help to facilitate the reproductive cycle of angiosperms.
Each flower part has a specific function.

Labelled Flower

A labelled, bisected specimen of the Erigeron glaucus, more commonly known as the Daisy. The reproductive (carpel, stamen, anther, and sepals) and non-reproductive structures (receptacle and pedicel) of the flower are displayed. Photo by Nick White.

Pedicel: The stalk of the flower

Receptacle: The part of the stalk where the various parts of the flower are attached

Sepal: Acts as the base for the flower

Petal: Aids in attracting pollinators

Stamen: The male part of a flower

Anther: The part of the stamen where pollen (male gametophytes) is made

Carpel: Houses female gametophytes

20150305_153229

Example of the most commonly cultivated fruit, the citrus fruit of a Rutaceae, commonly called an orange. Photo by Nick White.

After fertilization, the ovule transforms into a seed, and it is surrounding tissues evolve into a fleshy fruit. The fruit protects the seed and also promotes it’s dispersal to a wide geographic range. Much like flowers, fruit also has a large diversity among species. Some is meant to be dispersed by the wind, but many rely on animals to disperse it. Whether by having hooks to hook on to an animal’s skin or fur or being sweet and nutrient rich to promote being eaten, digested, and fertilized by the animals that carry them off (Carter, 1997).

 


 References


Angiosperms. (2008). In L. Lerner & B. Lerner (Eds.), The Gale Encyclopedia of Science (4th ed., Vol. 1, p. 217). Detroit: Gale.Carter, J. (2014, January 17). Angiosperms. Retrieved March 6, 2015, from http://biology.clc.uc.edu/courses/bio106/angio.htm

 Dilcher, D. (2000). Toward a new synthesis: Major evolutionary trends in the angiosperm fossil record. Proceedings of the National Academy of Sciences, 7030-7036. Retrieved March 6, 2015, from http://www.pnas.org/lens/pnas/97/13/7030#info

Hedges, S., & Kumar, S. (2009). Plants. In The Timetree of Life (pp. 133-137, 162-165). Oxford: Oxford University Press.

Soltis, D., Soltis, P., & Edwards, C. (2005, June 3). Angiosperms: Flowering Plants. Retrieved March 6, 2015, from http://tolweb.org/Angiosperms/20646/2005.06.03