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 1. Details of Module and its Structure

Module Detail

Subject Name <BOTANY>

Paper Name <Plant Physiology II>
Module Name/Title <Biomass allocation and Allometry>
Module Id
Pre-requisites Basic knowledge about biomass allocation and allometry, effect
of environmental factors and ontogeny on biomass allocation
pattern
Objectives To make the students aware of the components of biomass
allocation and allometric patterns.
Keywords Biomass, shoot/root ratio, root, stem, and leaf mass fraction,
allocation, partitioning,

Structure of Module / Syllabus of a module (Define Topic / Sub-topic of module )

<Biomass allocation <Sub-topic Name1>, <Sub-topic Name2>

and Allometry>

2. Development Team

Role Name Affiliation

Subject Coordinator <Dr. Sujata Bhargava> Savitribai Phule

Pune University
Paper Coordinator <Dr. Sujata Bhargava>

Content Writer/Author (CW) <Dr. Dhiraj Naik > Janakidevi Bajaj

Science College
Wardha
Content Reviewer (CR) <Dr. Sujata Bhargava>
Language Editor (LE) <Dr. Sujata Bhargava>

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 TABLE OF CONTENTS (for textual content)
1. Introduction
2. Methods of quantifying biomass allocation
3. Effect of environmental factors on biomass allocation
4. Effect of ontogeny on biomass allocation
5. Effect of competition and plant habit (herbaceous versus woody plants) on biomass allocation
6. Importance of allometry in forest and vascular plant productivity

Introduction
Plants fix newly synthesized carbon in leaves in the form of carbohydrates and transfer newly fixed
carbohydrates to different organs. In natural conditions, plant constantly sense the surrounding
environment and allocate nutrient and photosynthetic intermediates in different organs. In
natural conditions, biomass and resource allocation in plants is the central concept in modern
ecology, which provides a strong basis for different strategies of plants to respond and survive
under different environmental conditions. Allocation in plants is simply defined as a ratio of driven
process (e.g. shoot : root ratio, leaf : root ratio) and proportional (or fractional) process. It is also
called as partitioning. During the complete life cycle of plants, they tend to obtain range of sizes,
which is primarily driven by amount of resources available to the plants to grow.
Size is an important indicator of overall fitness and subsequent survival in natural conditions. Large
plants tend to have higher absolute fecundity (i.e. reproductive success) than small plants. Large
plants are also found to be better competitors than small plants. The ontogenetic age (i.e.
meristem age) also affects plant sizes. During early stage of plant’s life, (e.g. seedling stage after
seed germination), plants are smaller in size, whereas old plants tend to be larger. Therefore, plant
size is driven by various factors, majority of which includes age and resource (sometimes, referred
as nutrient) availability. This size dependent growth allocation is called as allometry. Allometry is
quantitative relationship between growth and allocation. Plants being static living organism
experience various growth phases. These growth phases respond to various biotic and abiotic
stresses and therefore they show change in an allometric trajectory (slow changes in growth phase
with reference to time) in response to the environments. This phenomenon is plasticity in

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 allocation. Being plastic in nature, plants continuously show changes in growth and allocation
pattern.
In this module, we shall study methods of quantifying biomass allocation, factors affecting
biomass allocation, ontogenetic effect on allocation and allometry.

Methods of quantifying biomass allocation

Biomass allocation is the realized distribution of dried biomass over the various plant organs. A
rigorous analysis of biomass allocation is required for the performance evaluation of plants
experiencing vastly different environmental conditions or in the growth comparison of different
species or different genotypes. Three methods are in common use for analyzing biomass allocation
patterns. These are root:shoot ratio, biomass fraction and allometry.

1) Allocation approach- shoot:root (S:R) ratio:

The first method is shoot:root ratio, also known as ratio allocation approach or biomass ratios.
This biomass ratio method is widely used in plant growth analysis. Mathematically, it is expressed
as:

Shoot: Root ratio = Shoot dry weight (in g)/ Root dry weight (in g) Eq.1

Percent Shoot: Root ratio= [Shoot dry weight (in g)/ Root dry weight (in g)] ×100 Eq.2

In many plant growth studies, the shoot:root ratios are reported as descriptors of the above-
versus belowground balance of biomass and thus they are indicative of plant responses to growing
conditions. There are few limitations of this measure. The ratio changes as plant grow. In
herbaceous plants, shoot: root ratios typically increase with age and size due to continuous
investment of fixed carbon in above-ground structures. Trees in a plantation forest would also
show a progressive reduction in root : shoot ratio, and especially after canopy closure where a
steady increase in stem biomass contrasts with biomass turnover of canopy and roots and thus
predominates in determining root : shoot ratio. S:R ratio indicates the trade-off between above-

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 and below- ground biomass. For example, in experiment with Deschampsia flexuosa, the S:R ratio
is constant over time in high nutrient treatments, but steadily decreases in nutrient deprived
conditions, suggesting plastic adjustment to increase below-ground resource capture.

Figure 1. Biomass allocation as affected by nutrient availability. Data are shown from an
experiment with Deschampsia flexuosa as described by Poorter et al. (1995). Data for high-
nutrient plants are given in black ,for
low-nutrient plants in red. (A)
Shoot:root ratio over time (B) Leaf
Mass Fraction (LMF)values (C) Leaf
mass(LM)plotted against stem plus
root mass (SMCRM) of plant (D) Leaf
mass plotted against stem plus root
mass for all individual plants (E) Leaf
mass plotted against total plant mass
(TM)for low-nutrient and high-
nutrient plants. (F) Leaf mass plotted
against total plant mass throughout
the experiment. (Adapted from:
Poorter and Sack (2012), Frontiers in
Plant Sci.3 (259):1-10.

Although, S:R ratios are simple to represent, this method has various pitfalls and disadvantage. a)
Typically ratios are nonnormally distributed. In literature, this statistical issue (i.e. normality of
data) has been often neglected. These S:R ratios should be log transformed prior to any statistical
analysis along with averages.
b) S:R ratios have conceptual drawbacks. It is unsatisfactory to describe biomass allocation in two
compartments. The combination of stem and leaves into one compartment as shoot does not
acknowledge different functions these organs have. Korner (1994) has given a realistic data on this

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 point. They compared biomass allocation of full grown trees of evergreen coniferous and
deciduous species. In this experiment, S:R ratio values were higher for deciduous species, and in
comparison with tree seedling and herbaceous plants, S:R values of both types of tree did not
show any variation. However, analyzing the same data in three compartments such as leaf, stem
and roots, then it appeared that biomass allocation to leaves was 3- fold higher for conifer species.
Also they found significantly huge difference between mature trees and herbaceous plants. This
concludes that allocation approach using biomass fraction (in three compartment- leaf, root and
stem) are far better estimate of biomass allocation as compared to S:R ratios.

2) Allocation approach- biomass fraction

Allocation approach can be characterized by fractions, expressing the biomass of each organ
relative to that of the total plant. Mathematically, it is expressed as:

Leaf Mass Fraction (LMF) = total leaf weight (in g) / total plant weight (in g)
Eq.3

Stem Mass Fraction (SMF) = total stem weight (in g) / total plant weight (in g)
Eq.4

Root Mass Fraction (RMF) = total root weight (in g) / total plant weight (in g)
Eq.5

The term ‘fraction’ rather than the more commonly used term ‘ratio’ is preferred, as it is the sum
of all fractions has to add up to 1 or 100 (if it is reported in percent), thereby providing an easy-to-
understand scaling in the dataset. The advantage of the use of biomass fractions over S:R values
are that LMF, SMF and RMF form an integral part of the concept of growth analyses and carbon
economy (Poorter and Pothmann 1992). Another advantage is that biomass fractions are less
sensitive to small changes in allocation than are S:R values, especially when roots form less than
20% of the biomass. These biomass fractions (LMF, SMF and RMF) can distinguish changes in

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 allocation over the time and with given environmental or growth conditions. In the case of
Deschampsia flexuosa, the LMFs lightly decreased over time for the high nutrient grown plants,
where as the decrease was much stronger in the low nutrient grown plants (Figure1B). At the final
harvest, LMF was 0.46 for the high-nutrient plants, and 0.31 for the low-nutrient plants,
representing a substantial and statistically highly significant (P <0.001) difference in allocation to
leaves. Although, as compared to S:R ratios, biomass fractions represents better in terms of
growth, this method has various pitfalls and disadvantage. a) Similar to ratio values, biomass
fractions are not normally distributed. The most appropriate methods to transform biomass
fraction data are arcsine and logit transformation. b) Differences in biomass fractions can
confound ontogenetic effects with treatment and species differences. Biomass fractions
misrepresent changes in biomass allocation, appearing as if due to treatment effect, which may
not be the case in if biomass fraction data represent active growth and developmental phase of
plants.

3) Allocation approach- Allometry

The main alternative to biomass ratios and biomass fractions is the analysis of allocation using
allometry, a size-dependent growth relationship. Allometric analysis is ideal method to analyze
which of the organs changes its growth response relative to the others, and at what size this
occurs. Thus, the allometric analysis is specifically able to avoid the problems described in figure
1F, because this analysis does not include an additional calculation (For example, in case of ratios
and fractions, the one organ mass is divided by others or total plant mass, which represent simple
mathematics). It is also not affected by the variability of having a numerator and denominator, a
down side of using ratios described in figure 1E.
The term ‘allometry’ is represented by the power function. Mathematically, it is expressed as:
Y=bXa Eq.6
where Y denotes the size of a plant organ (as gauged by its weight), X is a comparable measure of
the size of the plant minus the size of the plant organ of interest (e.g. root, stem and leaves), b is
the allometric constant, and a is the allometric (scaling) exponent. The formula to calculate
allometric relation is typically expressed logarithmically as:

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 log Y = log b + a × log X Eq.7

In this form, we see that log b is the Y-intercept (the numerical value of log Y when log X=1) and a
is the slope of log Y versus log X (Niklas 2004). Allometric relationship provides exact relationship
between any two plant organs or compartments without interference from changes in other
compartments.

Effect of environmental factors on biomass allocation:

Plants respond to environmental conditions spontaneously. Plant growth, development and
ultimately survival depend upon environmental variables such as light intensity, water availability,
temperature, which they face during the life time of plant. These factors strongly affect biomass
allocation pattern.

Effect of Light and elevated carbon dioxide concentration-

Under low light conditions, plants photosynthesize slowly and fix small amount of carbon. Under
the shade or low light conditions, they require less amount of nutrient due to their slower growth
and need less water due to decreased stomatal conductance. Shade loving plants allocate more to
leaves and stem as compared to roots as compared to sun loving plants. This observation is
referred to in the literature as ‘functional equilibrium hypothesis’. That is plants will allocate more
biomass to leaves if the limiting factor for growth is aboveground resources such as light. This is
observed when S:R ratios were plotted against daily light quantum inputs. Under lower range of
daily quantum inputs, S:R ratios were high. However, when same data was represented in the
form LMF, RMF and SMF, the biomass fractions did not comply with ‘functional equilibrium’
concept. More often, under varying light intensities, constant LMF was reported in various plant
species (Figure 2a, 2b, and 2c). In increased CO2 concentration treatments, plants grown at
elevated CO2 did not change their allocation at all (Figure 2d). However, herbaceous species and
woody species differ in response to elevated CO2. Herbaceous species showed increase in S:R
ratios whereas woody species showed decreased in S:R values.
Effect of nutrient and water-

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 Nutrients and
drought are the most
complicated factors
for biomass allocation
studies.
Figure 2. Dose–
response curves of
the absolute response
of the fraction of
whole plant mass
represented by leaves
(LMF; red line), stems
(SMF; brown line) and
roots (RMF; blue line)
to 12 environmental
factors: (a) daily
photon irradiance
(DPI); (b) red : far-red
(R/FR) ratio; (c) UV-B;
(d) CO2
concentration; (e)
ozone; (f) nutrient availability; (g) water availability (drought stress); (h) waterlogging; (i)
submergence; (j) temperature; (k) salinity; (l) soil compaction. Data are a compilation of the
available literature. For each environmental factor, a reference condition was chosen, and data for
each species in each experiment were subtracted from the allocation values observed or
interpolated for that reference level. (Adapted from: Poorter et al., (2012),New Phytologist 193
(1):30-50).

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 The response to drought is noteworthy, because plants subjected to mild water stress are often
very nearly the same size as control plants, with only a small increase in RMF (Figure 2g). Only
when plants are subjected to severe drought – that is when biomass is reduced by > 50% of that of
control plants – is a strong increase in RMF found. Changing allocation too quickly might then
result in a suboptimal growth after restoration of the water supply. Interestingly, LMF hardly
changes over the entire trajectory of the drought dose–response curve. Waterlogged plants show
a decrease in RMF (Figure 2h), which may partly be a reflection of dying root tissue. The reduced
RMF could contribute to the reduced stomatal conductance observed during waterlogging
experiments in intolerant species. There was a difference here between woody species, which
increased LMF on waterlogging, and herbaceous species, which did not (P < 0.05). Full
submergence of land plants is a special case, as they will not gain in biomass after the onset of this
treatment. A number of specialist species can survive if they are able to restore contact with the
air. To this end, they strongly elongate, partly by re-allocating biomass (especially starch) to stems
and petioles (Figure 2i). The change in RMF, however, is not stronger in species generally found in
submergence-prone habitats.

Effect of Salinity temperature and mechanical stresses (wind, soil compaction)-

Low temperatures decrease SMF, LMF and increase RMF (Figure 2j). A range of plant functions are
impaired by low temperature (photosynthesis, nutrient uptake, growth), but reduced rates of
water uptake are a probable cause of increased allocation to roots. Salinity negatively affects both
photosynthesis and transpiration, and minimally affects allocation (Figure 2k). Soil compaction has
a strong effect on allocation, but only at severe levels, in which case RMF increases strongly
(Figure 2l). Other abiotic factors have also been reported to affect biomass allocation. For
example, RMF generally increases and SMF decreases with wind exposure, stem flexure and other
mechanical effects on shoots.

Effect of ontogeny on biomass allocation

One of the main sources of plant size and plant biomass variation is age. An alternative but precise
term to define growth pattern is ontogeny. Ontogeny of seed plants takes place in distinct phases:

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 embryogenesis, seed formation, seed germination, vegetative development, reproductive
development, senescence and death of mother plant. As plant grows over the time, they readjust
morphology, allocation to different organs and leaf, stem and root physiology. This is adjustment
in other words called as ontogenetic drift. An adjustment in biomass allocation changes a response
to resource variability (i.e. nutrient, light and growth space availability) or as ontogenetic drift or
both. However, it is difficult to distinguish the variation in biomass allocation due to
environmental gradients versus ontogenetic drift. Ontogenetic drift is the change of a biological
trait in such a predictable way that it can be presented as a function of plant growth or
development. Biomass allocations to leaves, stems and roots have been intensively investigated
from this perspective. For annual plants, it was found that the proportion of allocation to roots
declines during their development, but for perennials, relative biomass allocation to belowground
(i.e. roots) was proposed to increase as they grow up. Therefore, some studies have concluded
that biomass allocation pattern to different organs is size-dependent, i.e., results from ontogenetic
drift.
Other variables of biomass allocation, such as leaf dry mass per unit leaf area (MA), fraction of
total plant dry mass in leaves (FL) and leaf area ratio (total leaf area per unit total plant mass; LAR
= FL/MA) have been identified as the major structural traits determining the potential for light
harvesting and accordingly species status in the community (Poorter and Nagel, 2000). In trees,
MA strongly increases with increasing tree height and age due to a juvenile to mature phase
change and possibly also due to decreases in water availability with increasing tree size (Niklas
2014). Age-dependent increase in MA and decrease in LAR have also been observed in the
perennial herb Leontodon hispidus (Niinemets, 2004), suggesting that the age-dependent trend of
increasing MA, that results in decreased light intercepting leaf surface area, is more general across
different plant functional types. Because many plant qualitative traits, e.g. formation of taproot,
stem and flowers depend on specific plant ontogenetic status rather than on absolute age, plant
age was characterized by a relative ten-level age scale (Weiner, 2004). Although the age
dependent changes in plant traits have been associated with modifications in accumulation of
total plant mass with calendar age (Poorter et al.,2012), ontogenetic changes in plant structure
and biomass allocation also occur at a common plant size (Weiner, 2004).

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 Effect of competition and plant habit (herbaceous versus woody plants) on biomass allocation
In natural conditions, plant species grows in competitive environment. Neighboring plants
influences biomass allocation which depends upon the plant density as well as interspecies and
intraspecies status of that plant. Research findings of biomass allocation with competition stress
studies shows that the average response to increasing plant density was a small decrease in LMF
and RMF and increased in SMF. An increased in RMF with increasing plant density was observed
only when plants in competition experienced low nutrient resources. In highly densed plantation,
plants increase the length per unit stem mass. One of the most important factor for competition
induced stem length increase in R/Fr ratio sensed by the phytochrome system. Increased stem
allocation could be considered as adaptive response of the shade avoidance syndrome. Patterns of
biomass allocation are important determinants of competitive interactions between plants. The
relationship between a plant's biomass, height and projected leaf area determines the efficiency
with which plants are able to intercept and absorb light for photosynthesis. In dense stands of
vegetation, light availability decreases with increasing depth in the canopy. In such stands, taller
plants will have an advantage because they are able to project their leaves in the highest positions
of the canopy where they receive the highest light intensities (photosynthetic photon flux density;
PPFD) while shorter plants grow in the shade of their taller neighbours (Weiner, 2004). On the
other hand, there are costs related to increased height: plants to invest a disproportionate
amount of biomass in support tissue (i.e. stems), and leaf (i.e. leaf laminae) mass per unit of total
above-ground mass (LMR) is generally found to decrease with increasing plant height. In addition,
leaf area per unit leaf mass (specific leaf area; SLA) is often found to decrease with increasing
PPFD. Thus, leaf area per unit above-ground mass (leaf area ratio; LAR), which is the product of
LMR and SLA, can be expected to decrease with increasing plant height and light availability per
unit leaf area.
Plant habit influences biomass allocation to leaves, stem and roots. A substantial difference is
expected between herbaceous and woody plant species, particularly substantial investment in
woody organ i.e. stems in forest tree and woody shrubs. Indeed, herbs in the vegetative phase of
the growth have an LMF of 0.5, whereas the LMF of mature trees is approximately 0.03. Within

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 the group of woody species, there is strong phenological contrast between deciduous and
evergreen species. If plants of higher leaf longevity are associated with infertile and ⁄ or dry sites,
we may expect evergreens to have higher allocation to roots and lower allocation to leaves. Thus,
the higher LMF may simply be the consequence of leaf longevity rather than higher assimilate
partitioning per se. In this way, conifers are able to compensate for their lower physiological
activity per gram of leaf. The majority of comparative experiments published are restricted to
either monocotyledonous or eudicotyledonous species. However, for the experiments that
comprised both, a higher LMF was found for eudicots, paralleling the allometric analysis.

Importance of allometry in forest tree productivity

The allometry studies the relative size of organs or parts of plants. The use of allometry is
widespread in forestry and forest ecology. The study of allometry is extremely important in
establishing the forestry practices in relation to carbon budgeting and carbon sequestration
analysis. The most common methods to assess the biomass is primarily based on forest tree
allometric equations. The tree allometric equations are helpful for evaluating forest tree volume,
bioenergy and carbon cycling. Different methods exist for developing tree allometric equations
depending on the objective (commercial volume, bio-energy, biomass or carbon), forest type
(mono-specific plantation or pluri specific forest), tree size, accessibility of the tree, forestry law,
technical, financial and human capacities (Picard et al. 2012). As a consequence, the quality of the
biomass estimates varies among allometric equations and depends on how the allometry has been
constructed.
Figure 3. Diagrammatic representation of
forest tree for allometric analysis.

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 Allometric equations based on tree volume and dried biomass is relatively difficult, time
consuming and requires heavy logistics and manpower. Destructive tree harvesting is sometimes
not possible based on forest site and location and also depends on forest law. Nondestructive
techniques are available for quantifying tree biomass using allometric equations. Easily available
measurements such as diameter at breast height (DBH) and tree height are enough to build
allometric equations (Figure 3). The general allometric equation is Y = β Xα where "Y" is a biological
variable (such as tree height or DBH), "β" is a proportionality coefficient, "α" is the scaling
exponent (which is equal to the slope of the line when plotted on logarithmic coordinates), and
"X" is some physical measure such as body volume or body mass(M). While α is often quite similar
between very diverse organisms, β differs from species to species.

References:
1. Bazzaz, F. & Grace, J. Plant resource allocation. (Academic Press, 1997).
2. Chave, J. et al. Tree allometry and improved estimation of carbon stocks and balance in tropical
forests. Oecologia 145, 87–99 (2005).
3. Enquist, B. J. Universal scaling in tree and vascular plant allometry: toward a general
quantitative theory linking plant form and function from cells to ecosystems. Tree Physiol 22,
1045–1064 (2002).
3. Körner C (1994) Biomass fractionation in plants: a reconsideration of definitions based on plant
functions. In ‘A whole plant perspective on carbon–nitrogen interactions’. (Eds J Roy and E
Garnier) pp. 173–185. (SPB Academic Publishing: The Hague)
4. Niklas, K. J. Plant allometry: is there a grand unifying theory? Biol Rev Camb Philos Soc 79, 871–
889 (2004).
5. Poorter H, Pothmann P (1992) Growth and carbon economy of a fast growing and a slow-
growing grass species as dependent on ontogeny. New Phytologist 120, 159–166.
6. Poorter, H. et al. Biomass allocation to leaves, stems and roots: meta-analyses of interspecific
variation and environmental control. New Phytologist 193, 30–50 (2012).
7. Poorter, H. & Sack, L. Pitfalls and possibilities in the analysis of biomass allocation patterns in
plants. Frontiers in Plant Science 3, (2012).
8. Weiner, J. Allocation, plasticity and allometry in plants. Perspectives in Plant Ecology, Evolution
and Systematics 6, 207–215 (2004).

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