Modification of The phenotype

Plasticity of the phenotype

Much of the phenotype variation encountered by the taxonomist is the
result of the plastic respone of the individual to factor of the environment. Such
phenotype plasticity has tened to be underestimated in taxonomic and
evolutioanry work in plants. From a practical point of view the problems it poses
are most acute in certain groups such as aquatic and marsh plants and in certain
regions such as the tropics, where van Steenis notes that using it for taxonomic
delimitation has given rise to a multitude of paper species and varieties without
any taxonomical value.
Development of the individual follows a more or less orderly pattern and
sequence to produce an organism of specific structure.These developmental
activities are nowadays explained in terms of gene action and the production of
specific growth substances, and come within the province of the developmental
physiology and morphogenesis.
An important differnce between plants and all but the smallest animals is
that growth is restricted to certain parts of the plant body. These are permanently
meristematic regions-regions of inderterminate growth, resulting in the continuous
formation and development of organs. Such regions provide good and extensive
material for morphogenetic studies.
This difference is related to the sedentary life of plants, with large surface
areas providing for nutrition. The stationary habit also makes individual plants
more susceptible to environmental fluctuations, and their plasticity provides
morphogeneticist with the possibility of studing the effect of the environmental
factors and presents taxonomists with numerous problems.
Although the plant body as a whole is relatively variable and not subject to
the same determinism as in animals, the varios organs from a clearly organised
whole. Individual organs such as leaves, carpels,etc. Have a much more precise
from than , say, the number of branches of a tree, but it is quite evident that the
overall shape of a tree, despite the variation in the number and position and
relative length,etc. Of the axes, is clearly recognisable and characteristic for each

species. There is in fact some kind of homeostasis or regulatory proses wich keeps
development within certain norms.
The number of organs, branching and sixe , tend to be more easily
modified in development than form and pattern. This is largely due to the period
od growth of the meristem and the degree of cell elongation at a later stage of
growth. These faetures are easily modified by factors such as temperature, ater,
loght and nutrition as discussed belows.
Genotype and Phenotype
As we have seen, living plants, like all living systems, are today regarded
as organised systems wich maintain and adjust themselves, through their capacity
for homeostasis, to the environmental flux. A distinction is made between the
genotype wich remains unaltered (except under certain circumstances) and the
outward appearance or phenotype wich alters in structure in respone to changes in
the environmnet, these alterations sometimes being partially reversible. There
appears therefore to be a fixed factor the genotype and a variable expression the
phenotype in respone to varying environmental conditions, and a homeostatic
realtionship co-ordinating the system.
It is only recently that the dichotomy between heredity (genotype) and
environment, and the effect pf each on the phenotype, which has been
overstressed in the past, has been questined. The effects of heredity and
environmental factors always operate jointly and frequently it is not possible to
disentangle them. As Waddington has expressed it organism and environment
are not two separate things each having its own character in its own right, which
come together with as little relationship as a sieve to a shovelful of the pebbles. It
has been shown, on the contrary, that there is a close integration between the two :
the environment can modify the channel which the form of the organism can
take. Again, as Sinnott in his recent review of morphogenesis says, a visible trait
(i.e. morphological faeture) is the development reaction of a spesific (and
constant) genetic constitution to a specific environment.
The response of the different faetures of the plant to different
environmental conditions or factors considerably. Some characters, such as leaf

arrangement or floral structure, remain more or less unchanged under a range of

different conditions. These are frequently referred to in taxonomy as good
characters because of their relative constancy and are regarded as genetically
fixed. Other features such as leaf shape, stem height, time of flowering,etc.
Develop differntly under different environmental conditions. The term phenotype
plasticity is applied to the variations shown by the phenotype in response to
environmental fluctuations. The same genotype may, accoriding to the
environmental factors prevalent, give rise to different phenotype. The plasticity
may be so wide to overlie or even obsurce the genotype variability. In fact, the
distinction between genetically determined characters and environmentally
induced characters is a false one, since both types of characters are the result of
genetical inheritance and environmental control. In other words, each genotype
has a range of expression is narrow the plants are termed stenoplastic or broad the
plants being called euryplastic. It is the inherited response which produces the
character or phenotype,not just the environmental factors. The adaptive or nonadaptive nature of these differences is discussed below in connection with
ecotypes.
The phenotype can be regarded in the light of what we have just discussed
as the product of the interaction of all genes with each other and with the
environment. It is the phenotype which is exposed to natural selection, and the
phenotype that the taxonomist must be primarily concerned with, not the
individual genes, chromosomes or genotypes, for the latter obviously can only be
perceived through their phenotype effects.
The factors which affect the plasticity of the phenotype are claerly of great
concern to the taxonomist. It is, for instances, very necessary to find out if
variation is due to genetically inherited differences or to the particular range of
environmental factors which produce largely reversible modifications from the
same genotype. The fact that similarity in phenotype may be due either to
environmental modificarion or to genotypic differentiation makes such similarities
unsure guide of genetic reaktionship. They are usually disregarded by
evolutionists due to their possibly non-genetic nature. Taxonomic may be misled
by phenotype variation in their selection of taxonomic characters. It is not of

course possible to determine the narute of the variation whether genotypic or

environmentally induced in the hebarium or even in the field, and comparative
cultivation is normally necessary. It may be possible to argue in some cases by
analogy and extrapolate fron experience of other situations, but there are always
attendant dangers on such indirect methods. Nonetheless, taxonomists have
frequently to attempt it.
Growth Substances
The regulations of growth and organised development is largely achieved through
the agency of growth substances. Recent morphogenetic studies suggest that many
of the features of form and structure on which the taxonomist relies (such as
symmetry of flowers, inflorescensce structure, number of floral parts, nodal
anatomy,etc.) are probab;y determined by complex patterns of balance between
gene-controlled cellular states at primordia: the latter are activated or deactivated
through hormonal control causing the initiation of particular development
pathways. The precise mode of action of these systems is not yet known, but it is
suggested that profound alterations in the phenotype can be caused by small
change in auxin balance or concertration and the time and rate of diffusion of
these substances.
The determination of teh particular pathway along which a tissue will develop
probably takes place at a very early stage of growth in the primordium, but
whether the normal developmental pathway isn continued to its particular end
may depend on substances which enter the developing primordium from the apex.
In Cannabias sativa the effect of treatment with2,3,5-triiodo-benzonic acid(TIBA)
is to modify the pattern of emergence of the perianth members and stamens to
form flowers of a tbular form but without deflecting their developmental
pathways. TIBA simulates the effect of the genetical changes which, in the
course of evolution, have brought about such transitions as that from polypetaly to
sympetaly.
Major structural and functional alterations may be induced in intact flowers by
externally applied auxins, such as the suppression of the corolla and androecium
in species with diclinous flowers and the hypertrophy of the calyx and gynoecium

and the production of female or inter-sexual flowers in dioccious hemp. In the

latter case the ontogenetic development of the presumptive staments appears to
have been diverted towards the pathway producing carpels, the auxins having
possibly influenced some determining proscess in the flowers primordium.
Herslop Harrison concludes that since floral morphogenesis can be influenced
by applied auxins, we would expect that environmental factors which alter auxin
metabolism should have similar affect on the structure of flowers. That structural
alterations induced by environmental factors parallel those caused by auxins does
not necessarily mean that native auxins have been responsible, but the like lihood
is there.
Differences in auxin concentration at the apex of the plant can alter the form of
branching which may constitute an important taxonomic difference between
species, as in Asternovaeangliae and A. Multiflorus. The former has one main
stem whareas the latter is much branched, this has been shown to be correlated
with the amount of auxin present in the plant. The effect of growth substances on
flower induction and sex determination has been well studied and documented.