Shade-mediated phenotypic plasticity in the Synsepalum dulcificum and in

Ilex aquifolium

Light is a very important element in nature and pretty much all plants and flowers are
subjected to at a minimum level of light intensity as shade through their lifetime.
Different light gradients within plant canopy takes numerous stress factors, as changes
of leaf characteristics on the light intensity as well as the diminished R to FR ratio from
the pinnacle to the lower part of the leaves of the three branches, generally in the
course of leaf formation. Specific leaf area (SLA which represents the rapport between
leaf area and leaf dried out weight) is a morphological characteristic with higher
plasticity to light variations including the morphogenetic regulation of foliage that will
probably increase the surface of the leaf in partial sunlight to be able to capture much
more light while there's a physiological limitation to the entire leaf volume. Specific leaf
area (SLA ) reflects also leaf thickness as well as the relative ability of assimilation of
light, transport of nutrients and stereome.
The thickness is higher for the leaf’s exposed to full sunlight compared with the ones
that are exposed to partial sunlight and this aspect represents an important factor in
assimilating the light. Sun foliage have a better photosynthetic percent on a leaf area
basis which represents a greater chlorophyll percent, a much lower light harvesting
chlorophyll proteins percent, a low stacking number of thylakoids, along with a greater
nitrogen content material for each unit of leaf. Part of nitrogen is consumed in the
synthesis of proteins in plants. The scientist Hirose and his partner Werger describes
how “…optimal pattern of nitrogen distribution that maximizes the canopy
photosynthesis was determined. Actual distribution of leaf nitrogen in the canopy was
more uniform than the optimal one, but it realized over 20% more photosynthesis than
that under uniform distribution and 4.7% less photosynthesis than that under the
optimal distribution. Redeployment of leaf nitrogen to the top of the canopy with ageing
should be more effective in increasing total canopy photosynthesis in a stand with a
dense canopy than in a stand with an open canopy“ publicized in their article
“Maximizing daily canopy photosynthesis with respect to the leaf nitrogen allocation”
Changes in the characteristics of leaf to survive to reduced light raise the ability to
absorb light in the process of photosynthesis and in this way, the quantity of carbon
dioxide which is lost through respiration is diminished.
In order to use effectively the light for plants growth, a few aspects have to be taken in
consideration like the direction of light through the leaves of the plant, the cost of
radiant energy and the way that surface of leaf’s intercepts the solar energy. If all these
conditions are achieved, then a proper use of light for development of the plants is
reached. The shade tolerance is the least light to which a plant can stay alive. Shade
leaves and sun are actually more delicate in tolerant species, the light demanding
species are definitely more plastic compared to shade tolerant ones. The capacity to
adjust to different light environments can be essential to woody species since new
plants developed on the soil surface of the forests and they are exposed to strong
sunlight which comes through gaps formed in between the trees. These little species of
plants must defend themselves against the strong light using their photo inhibitors.
Strong sun exposure can trigger continual reduction on the photosynthetic capacity via
impairment of second photosystem reaction centers in the foliage of plants which have
encountered gap openings. Complete expression of plasticity in shade tolerant species
relies upon availability of additional resources as nutrients or water. The purpose of
adaptation of this species to shade is connected with different characteristics.
Even if general plasticity could be lower in shade tolerant species, other traits are able
to adapt to full sunlight than those which cannot survive in shade. The species of plants
that can adapt to different grades of shade, can adapt to different environments
because of traits they have on overall as a plant. Synsepalum dulcificum is described by
Enoch Gbenato Achigan-Dako in his article “Current knowledge and breeding
perspectives for the miracle plant Synsepalum dulcificum(Schum. et Thonn.) Daniell” as
“previously known as Bumelia dulcificum … or Sideroxylon dulcificum …, belongs to
Sapotaceae, a large plant family with circa thirty to forty genera about which knowledge
is still obscure. The genera Synsepalum is confined to tropical Africa and consists of
about thirty species.... In tropical West Africa, eight useful species were identified
including Synsepalum afzelii …, S. brevipes…, S. cerasiferum…, S. dulcificum…, S.
glycydorum…, S. revolutum…, S. stipulatum…. Out of those species, S. dulcificum is the
most widely distributed, covering Ghana, Benin, Nigeria, Cameroun and Congo”.
E.G. Achigan-Daku is describing Synsepalum dulcificum as an “evergreen shrub” which
is reaching four meters in high. The author also mentioned that “leaves are entire,
alternate, symmetrical, and obovate-lanceolate to broadly lanceolate and pointed at
apex, with 4.3-7.5 cm long and 3.1-3.8 cm in width. The length/width ratio is 2:1. The
apex itself is obtuse and is occasionally slightly acuminate. The adaxial surface of the
leaves is glabrous and the abaxial surface is pubescent”. The leaves of this plant
contain no glands and they are grouped around a very small petiole.
On the lower part which is facing down the leaf epidermis is highly covered with very
thick doubled layer of hair. Small , white flowers which are bisexual, solitary or in
clusters enrich these evergreen plants. For each sepal is distributed a petal and a
stamen, totally being four or five of them in the calyx. For the mature plant, the flower
has the ovary in the top part of the stigma and is covered by a heavily hairy wall. The
berries which are the fruits are oval turning red from green when are ripe.
This tropical species contains a special glycoprotein which is used as a sweetener and
because is coming from a natural source, can be used successfully as a sweetener
substitute specially for people suffering from diabetes. Because of this glycoprotein
which was called “miraculin”, the fruit (miracle berry) is a promising economic
alternative to sucrose (regular sugar). At present, S. dulcificum is used in cosmetics
and food, although it's most extensively utilized by the pharmaceutical industry. As any
good , miracle things there are some impediments that slows the economic procedures.
One of those is the fact that S. dulcificum is a notoriously slow growing species,
generally found in home gardens and less often in farms. This is a big gap in the
availability of these promising berries. Empirical observations have indicated that
seedlings growing in home, tree-based gardens, are significantly stronger than those
exposed to full sunlight on open farms. Researchers tried to find solutions to the
problem related with the large scale of harvesting S. dulcificum by conducting a series
of tests and experiments. These were focused on the exposure to light of this plant.
Environmental induction as a suitable method of inducing flowering in several tropical
horticultural crops owes one of the ideas exposed by researchers. Others indicated that
photoperiod, nutrient, availability, and water happened to be the major environmental
factors that will result in flowering in woody species. In their study, Ohishi and
Yamazaki indicated that light exposure consistently accelerated flowering in the
blueberry, while in the white birch a well-balanced fertilization containing nitrogen,
phosphorus and potassium can help to promote the transition from the juvenile level to
maturity, but in addition to elevate flower production. In the case of S. dulcificum,
watering as well as inorganic fertilization have before been found to accelerate and
enhance flowering.
A related study was conducted by Valladares, F. together with another scientists, on
shade tolerance, plastic phenotypic response and sensitivity to photoinhibition in holly.
His data and research conclusions were published in “Shade tolerance, photoinhibition
sensitivity and phenotypic plasticity of Ilex aquifolium in continental Mediterranean
sites” article Ilex aquifolium L. is a plant that is found mostly in temperate forests and
also woodlands. This species of plant is very sensitive to drought that’s why the
regrowth of the plant is based mostly on seedling. In conditions of high humidity, holly
can develop in the sun or in the shade without no problems even though the plant
prefer partial shade. In dry regions where is hot and arid , holly will grow only in shade.
Holly is sensitive to low temperatures , so areas with these kinds of climate where is
colder, are not favorable to its growth. In temperate oceanic climate , this plant is at
risk to be replaced by other species because of increase carbon dioxide gas.
For their experiment, Valladares, F. and his team considered only one major aspect of
conditions for optimal growing of holly, which was the exposure to sunlight, direct or
indirect ( shade). The area of research was the temperate climate of Europe where
aside of high solar exposure are other stress factors like drought and high
temperatures. Valladares, F. et al. focused on “ characterizing the light environment of
seedlings and adults, and on estimating phenotypic plasticity for a range of
morphological and physiological variables. Phenotypic plasticity in response to variation
in resource availability is central to the environmental adaptations of sessile organisms
exposed to changing environmental conditions …, and plant species have been shown
to exhibit significant differences in their plastic response to light”.
A few hypotheses were followed through the testing, one of them implying that holly
should suffer and develop slow and hard in condition of a harsh climate as the
continental, Mediterranean. Also, holly which develops in Mediterranean areas should
be very sensitive to photo inhibitors in a high solar exposure together with other climate
stresses. The opposite to this hypothesis, the next one implies that holly should show
higher phenotypic plasticity in shade because it’s a shade tolerant species. In order to
test these hypotheses, they placed the seedlings and adult plants to shade and also in
high light and the plastic phenotype in both conditions of both stages of the holly. The
seedlings were exposed to different grades of shade and the plastic phenotype was
recorded as well as adult plants were exposed to shade and sun with specific
consequences on photoinhibition and water relation in the field.
In the following, a shade tolerance and phenotypic plasticity presentation will be made
on Synsepalum dulcificum (B) and Ilex aquifolium (holly)(A) together with the center
point within the seedlings section of the experiments, but not ignoring the other aspects
of these scientific studies (mineral nutrients addition for the "miracle" grow, and the
plastic reactions to light on adult plants of holly).
The experiment for Ilex aquifolium (holly) was taking place in Spain in three different
locations, all three locations being in continental Mediterranean climate characterized by
extended periods of drought in the summer and significant differences in temperature
from day to night or between seasons. It’s important to mention that these locations
were at an altitude of fifteen hundred meters approximately, so these locations were
actually forests which have an acidic soil and a very low capacity of retaining water and
not very rich in nutrients. Each forest had different characteristics, the main one being
the species of threes that are forming that forest. One of the forests had a very high
density which was formed by a high number of different species of threes, and the
other two had a different composition of three species which allowed more light to
come through it. Holly seedlings were collected at cotyledon stage, placed in pots with
soil from the forests, taken into the green house where they been watered every other
day. After two weeks they been placed into the experimental areas and watered to the
full capacity of the soil every other day. After five months of growth, the total growth of
plants was measured as well as their components including the leaves and total
structure and aspect of it was recorded. The test was repeated in the following year in
the same manner, but the experimental location was changed with others which had
more exposure to light. At the end of the second year the content of chlorophyll and
carotenoid was measured as well as highest rates of photosynthetic electron transport.
Holly seedlings have been removed from the ground and splinted into their
components as leaves, roots, stems in order to be tested and analyzed. To keep the
leaves away from oxidation and desiccation, a special way of transportation was used
like the black bags that don’t allow the light to get through. After that, the whole
organic material was dried out for forty-eight hours at sixty-five degree Celsius and
weighed. All this information was utilized to calculate certain leaf location, root: shoot
ratio, leaf region ratio, and distant relative growth rate. The measured parameters
showed no substantial differences in both trials. The data collected from these
experiments showed that holly can tolerate moderate shade but not deep shade which
is around one percent. In a very dense forest with an eight percent of light, the holly
seedlings can survive but to develop properly they need at least ten percent of sunlight.
Holly had a poor performance in high light also which is not making it suitable for open
fields. So, the best performance of this plant is around twelve percent sunlight which
place it in bright or moderated shaded forest (understories) in continental
Mediterranean climate. Photoinhibition of holly was significant in highlight exposure in
the field. This affected the photosynthetic capacity of the plant, but besides of highlight
exposure, low nitrogen availability was also a major factor for photosynthesis. The plant
which have a low percent of nitrogen in their leaves can reduce the chances of the
species to adapt to full sunlight. But for this experiment no major differences in
photosynthesis were recorded between sun and shade population of adult plants and
this is because of limited nitrogen availability. As a conclusion to Valladares, F. and his
colleagues’ experiments Ilex aquifolium (holly) can be considered an optional shade
tolerant plant in humid and moderate climates and obligate shade tolerant plant in dry,
continental areas like continental Mediterranean one where the experiment was
conducted.
Similar with Ilex aquifolium (holly) experiment was the one made by Dèdéou A.
Tchokponhoué and his team for Synsepalum dulcificum. Their experiment started in
the spring of 2015 and the scientific report called “. Shade and Nutrient-Mediated
Phenotypic Plasticity in the Miracle Plant Synsepalum Dulcificum” was publishes last
year (2019).
They used for the experiment specific seedlings which were obtained from mature, ripe
fruits. The fruits were peeled off and the seeds extracted. After that the seeds were
putted to germinate in certain conditions with a specific soil composition to make sure
that nutrients will provide good conditions for growth. This experiment took place in
Benin in Africa where the climate assured sufficiently humidity, moderate temperature (
around twenty-seven degrees Celsius) good quantity of rainfall and plenty of sunshine.
The objective of this experiment was to determine what are “ the effects of light
exposure and compost application of the survival, vegetative growth, reproductive
performance, biomass allocation, and plasticity of the transplanted S. dulcificum
seedlings”. The main factor tested was the light exposure and the seedlings obtained in
2015 after one month of germination of seeds that were mentioned above. Beginning
with the spring of 2016, on a period of fifteen months, part of the seedlings was
exposed to full sunlight and the other part were kept in moderate shade. The soil was
watered to fullest extent, but scientists wanted to determinate the best version of the
soil nutrition for the fastest growth of this plant because there wasn’t any previous
study made for the effects of organic nutrients in the soil for the growth of S.
dulcificum.
After the experiment was finally done and data collected some conclusions could be
obtained. Phenotypic plasticity refers to the ability of a plant to adapt to different
conditions. It was proven by the measurements made during the test that S. Dulcificum
showed a greater plasticity to different light intensity exposure compared with the
nutrients presents in the soil. Also, a darker green leaf, which indicates more chlorophyll
in the leaves was noticed on the plants kept in the shade compared with the ones
exposed to full sunlight.
The study showed the importance of light exposure of S. dulcificum. Besides of the
higher survival rate of seedlings under shade conditions, the period until first fruiting
was reduced which means an increase of proportion of seedlings entering the
reproductive phase. Another aspect that showed the benefits of the shade is the
vegetative and growth traits of this plant, the high of seedlings being double compared
with the ones from full sunlight exposure and also the size of the leaves were bigger.
Valladares and Niinemets indicated in their article “Shade tolerance, a key plant feature
of complex nature and consequences “ that “ in intermediate shade-tolerant species,
stem elongation was often found to be the most plastic trait.” The scientists who made
this experiment also found that the seedlings height represents the important plastic
trait in Synsepalum dulcificum which supported their hypothesis that this plant is an
intermediate shade-tolerant species. Once established the favorable conditions for S.
dulcificum which are reduced light exposure and rich nutrients soil, a promising
potential for growth and expansion in new environments for this species is taken into
consideration. These characteristics indicate a successful adaptive plasticity of
Synsepalum dulcificum, and give economic possibilities for future income generation,
stronger asset for smallholder’s farmers and overall economic growth of some
countries.

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