Lorenz G.

Palec 27 April 2019

SFI 271 Advanced Silvics

Journal Critique:
Growth and physiological response of six Australian rainforest tree species to a light gradient
J. Kelly, S. Jose, J. Doland Nichols and M. Bristow
Summary

Forest degradation is prevalent globally, and Australia is not an exemption due to land-
use conversion which is mainly caused by the population’s need for resources. The eastern
coast of the country, however, remains as fragmented rainforest. Large clearing/cutting
activities was dominant as the past land use conversion to agricultural lands, grazing lands and
timber lands occurred. In effect, the Australian landscape is now highly lacking the presence of
intact rainforests. Although forest restoration efforts have been recorded, this was primarily
missing on the information on rainforest species’ ecophysiological characteristics, resulting to
failure in the whole approach. Establishing the knowledge on both physiological and growth
responses of tree species to varying light regimes in applied natural sciences (i.e. forestry,
agriculture, horticulture) leads to success in the practice of natural resource management, like
forest restoration activities which aims to initialize forest succession where it is absent.
Knowing how to manage the restoration based on trees’ and plants’ inherent capability to
adapt to resource-limited conditions is nonetheless essential.
The authors found it vital to have baseline information of species responses to different
environmental conditions (in this case - light). The article focused on examining six Australian
rainforest species’ light requirements for facilitation of growth, morphology and photosynthesis
under natural and light-moderated conditions. This also specifically aims to: a) know how the
species adapts its photosynthetic processes with response to different light regimes; and b)
examine if the varying light regimes affect the species morphology, growth and biomass
allocation.
 Three successional stages (early, late and mature) were considered in selecting the six
seedling species to be used in the experiment: a) Cryptocarya erythroxylon (Ce); b) Elaeocarpus
grandis (Eg); c) Flindersia brayleyana (Fb); d) Flindersia schottiana (Fs); e) Gmelina leichhardtii
(Gl); and f) Heriteria trifolialatum (Ht). The study was conducted at the Southern Cross
University in Lismore, New South Wales, Australia. The experimental design used was a
randomized complete block design with three greenhouses as blocks, three tables within each
greenhouse corresponding to the three light regime treatments (10%, 30% and 60% full
sunlight). All seedlings were transferred from small pots from area nurseries into standardized
3.8L self-draining pots with same potting medium used. These were then let into open sunlight
for 15 days before being placed randomly into each treatment and into the greenhouse tables.
Field capacity (moisture) was maintained by watering for all seedlings, including standardized
fertilization (with Aquasol fertilizer mix). Rate of fertilization was determined by conferring with
knowledgeable nursery managers and using secondary data. For the data collection, light
response curves were generated using LICOR-6400 at different light levels (2000, 1500, 1000,
500, 200, 100, 50, 20 and 0 mol-2 s-1. Carbon dioxide pressure and temperature was
maintained constant across seedlings’ leaves during the measurement (for 10 randomly
selected seedlings per treatment). Relative growth rate (RGR) was computed as referenced to
Hunt’s (1978) method and formula. Leaf area was generated using LI-COR LI-3000. Specific leaf
area (SLA), leaf-mass ratio (LMR), stem-mass ratio (SMR), and root-mass ratio (RMR)
correspond to leaf display and biomass allocation and were computed by deriving the primary
data used in RGR calculations. Mitscherlich model equation was used to regress (non-linear) the
light curves generated, designating Amax as the light saturated photosynthesis, Aqe as the
quantum yield and LCP as the light compensation point. A two-way ANOVA was used in
determining significant morphological and photosynthetic responses of the seedlings. Tukey’s
multiple range test was employed if significant values were found to segregate means.
All species Amax increased from 10% to 30% full sunlight with additional differences in Eg
and Fb which are both early secondary species. At higher light levels, Eg, Fb, Ce, and Ht had
higher photosynthetic rates compared to the 10% treatment., although Gl had increased rates
in 10% full sunlight. Mean values for Aqe for all species except Fb were highest in 60% full
sunlight and decreases as light level declines. Most species’ LCP were significantly high in 60%
full sunlight treatment except for Fs. Aside from Ce and Ht, all species exhibited maximum stem
diameter in 60% full sunlight. On the other hand, height growth for all species was greatest in
60% full sunlight except for Fb and Gl. Highest RGR values were found for late and early
successional species, in addition, these also showed significant changes in SLA from low to high
light. Generally, LMR was highest in 60% full sunlight for all species while RMR results contrast
this as more resources were allocated to the roots in 10% sunlight.
It is known that light highly limits plant growth within rainforests as the dominant
canopy suppresses undergrowth and intermediate layers. How certain species adapt to this
situation determines their ability to compete for this precious resource. Knowledge on sun and
shade plant characteristics have also been previously studied by other authors. It was found, in
general, that trees have an increasing Amax up to a certain point where the rate declines in high
levels of light. The known photosynthetic response of shade-intolerant trees was shown by Eg
and Fb, as its sensitivity to lowering of light levels resulted to acclimation. On the other hand,
Amax of Gl (shade-tolerant) was relatively higher in low light levels. For the early stages of
growth, part of the results of this experiment may be able to explain how most rainforest
species are able to tolerate low light levels at the early stages of plant development and that
shade tolerance may be dependent on the morphological and carbon allocation capacity of
these species in shaded conditions. The results also suggest the general model for SLA, with the
species having higher SLA in low light conditions, although shade-tolerant species were
observed to be less plastic compare to the early and late successional ones. Seedlings grown in
low light levels had lower RGR compare to the individuals in high light levels. It is then
concluded in this study that the species which showed highest RGR were also the most plastic
(high SLA in low light levels). The maximum value for RMR in Ce, Fs and Gl were found in the
10% full sunlight, which may explain the low RGR since most of the resources were put into the
roots rather than used for leaf expansion. In addition, these species also showed high LMR in
high light treatments, which will also most likely decrease the resources allocated into the roots
for higher water uptake. Hence, varying responses to different light levels were found for all
species. Some were found to be most plastic and can be placed in both low and high light
conditions. All the late and mature successional species exhibited best growth in 30% or 60%
full sunlight. Highest sensitivity to light conditions was found for Eg and Fb.
Although shade house experiments cannot imitate the real light and moisture
conditions in the field, results have been relatively similar to field studies with respect to
species rank in growth parameters. Establishing a protocol by determining the ideal canopy
light levels for ideal plant growth may be beneficial for use in management of restoration
efforts. For practical application, Eg and Fb may be used as nurse trees in the field while the
other four species are recommended for the late stages of restoration. It should however be
observed that careful maintenance and species selection for species to be planted together
with Eg due to the species’ highly competitive root system and its ability for high growth rate in
high light conditions.

Insights/Comments:

 There must have been a mistake in the concluding part of the abstract as Eg and Fb showed
highest plasticity and acclimation to the different full sunlight treatments, which are mature
and late successional species, respectively, and do not represent most of the shade-tolerant
group.
 Although it is not in the scope of the study, I suggest that genetic characterization of
responsive species be studied to be more precise in determining the extent of acclimation.
 Other applied natural sciences which can use the results of this study are arboriculture and
agronomy.
 The objectives were well explained by using question-type format
 We are well aware of the terms shade-tolerant and shade-intolerant species. However, it
was not clearly defined and segregated in the species description of the ones used in this
study (early, late and mature successional species). These may be terms they use in
Australia. I may be much better to explain which of the species represent the two general
classifications.
 It was not stated whether the neutral density shade cloths were fabricated or what they
really where, or I suggest that if these bought commercially, it is better to describe the
specifications or brand to help other researchers conduct a similar experiment to other
species and possibly produce comparable results.
 It was not clearly stated how frequent watering was done (i.e. hourly, every 10 hours, daily,
weekly) in order to maintain field capacity.
 Although it was said that fertilization rates were discussed with nursery managers and
secondary information, it was not disclosed when the activity started, frequency of
application and when did it end.
 Although it was not really necessary in a publicized with respect to cost of publication, the
randomization method used for determining the 10 seedlings for photosynthetic
measurements were not stated.
 The specific light flux when the photosynthetic data was gathered was not defined. The
authors only stated partly cloudy to clear skies. It may have been better if they used a light
sensor/meter to determine existing light conditions.
 If there is a more recent formula for RGR computation than Hunt’s which was formulated in
1978, it is better to use it.
 The conclusions of this research may support that the particular silvicultural system to be
used in rehabilitation/restoration maybe species-dependent (Beadle & Sands, 2004). For
example, if the target species is shade-tolerant and is not plastic to variations in light levels
(like Gl), it will be better to use shelterwood method rather than clearcutting. Clearcutting
may then be done when the target trees are of early successional stage species (like Eg).
 The results will be useful in plantation management, such as in the conduct of light
environment manipulation by altering stocking density (Beadle & Sands, 2004) depending
on what successional stage the target species to be facilitated is.
 We rarely see this kind of experiment/study being published for the Philippines. The results
clearly describe and document basic information on Australian
reforestation/restoration/timber species responses to environmental conditions (in this
case, light levels). This kind of information maybe basic, but it is important for managing
 activities concerned with planting trees. As a forester, conducting such a research in our
country for our indigenous and endemic species maybe expensive and we may lack the
equipments (in terms of number and the equipment itself) needed to do this, but it will be a
great help in terms of species-level conservation in addition to successfully restoring our
degraded/idle lands and timber production.

References
Beadle, C., & Sands, R. (2004). Physiology and Silviculture. In J. Burley, J. Evans, & J. Youngquist,
Encyclopedia of Forest Sciences (pp. 1568-1577). Kidlington,Oxford: Elsevier Ltd.