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Box 5.4 Growth Analysis in Vriesea sanguinolenta (Stefan Wester, Cord Mikona, and Gerhard Zotz)

Box 5.4 Growth Analysis in Vriesea sanguinolenta (Stefan Wester, Cord Mikona, and Gerhard Zotz)

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Photosynthesis, Carbon Gain, and Growth


Box 5.4 (continued)


The area-based photosynthetic capacity of the youngest, fully expanded

leaf ranged from 2.2 to 13.4 μmol O2 mÀ2 sÀ1. It was significantly affected by

nutrient supply, light, and initial size (3-way ANOVA, p < 0.05), while on a

mass basis, only higher leaf nitrogen concentrations influenced photosynthetic capacity (3-way ANOVA, p < 0.05).

Relative Growth Rates

As in other epiphytes, growth in the studied plants was slow. Even under

the best combinations of water, nutrient, and light, the highest RGR did not

exceed 8 10À3 dayÀ1. All studied factors significantly influenced RGR, but

rarely independently of each other. For example, RGR was increased at high

light by more than 50 %, but only when nutrient supply was high. This

difference was consistently found in all size classes. Both improved water

and nutrient supply acted in a synergistic fashion, but their effects and

interaction were size dependent. Consistent with the results of an experiment

of shorter duration (Laube and Zotz 2003), the strongest stimulation of RGR

was observed in small individuals with an increase from 1.1 10À3 dayÀ1 under

low light/low water conditions to 5.7 10À3 dayÀ1 under high light/high water

conditions. SLA was consistently affected by light with an average reduction

of SLA in high light of 11 %.

The relative growth rate correlated closely with net assimilation rate

(NAR), which in turn was closely related to area-related photosynthetic

capacity. Specific leaf area, on the other hand, did not correlate with RGR

nor did LMR. The relationship of NAR and RGR differed among size classes

as shown in the following graph.



RGR, 10−3 d−1











NAR, g m−2 d−1

The slopes of the different size classes (symbol size and line thickness

indicate small, medium, and large plants) differ significantly, ANCOVA,

p < 0.05). In the smaller plants, a significantly steeper slope led to a higher



5 Physiological Ecology

Box 5.4 (continued)

RGR for a given NAR. This difference coincides with a significantly higher

LMR in smaller plants.


It is debated whether variation in RGR is primarily related to variation in

SLA (Lambers et al. 2008) or in NAR (Shipley 2006). The described results

support earlier conclusions based on an experiment of rather short duration

(Zotz and Asshoff 2010): NAR is the major factor associated with variation in

RGR in the studied tank bromeliads. The duration of our experiment (almost

1 year) was long enough to allow the production of an entire new set of leaves.

The fact that SLA varied significantly with light environment in all size classes

clearly indicates that the time frame was sufficient for possible changes in

morphological parameters to develop in these very slow-growing plants.


Atmospheric CO2, Net CO2 Uptake, and Growth

The concentration of CO2 in the atmosphere has continuously increased since the

onset of the industrial revolution and is expected to rise even more in future decades

(Solomon et al. 2007). The rate of photosynthetic CO2 uptake is not saturated in C3

plants under current conditions (K€orner 2003); hence epiphytes could actually

profit from this development because of improved water use efficiency. A few

studies have focused on the effects of elevated CO2 on epiphytes (e.g., Monteiro

et al. 2009; Raveh et al. 1995; Li et al. 2002a), but the interaction with other

important factors (e.g., water and nutrient supply) has rarely been analyzed. Studying CO2 in isolation may be problematic for an ecologically meaningful interpretation for the following reasons: (1) Water supply is assumed to be the most important

environmental factor for most epiphytes (Zotz and Hietz 2001) and (2) both water

and nutrient availability generally influence the effect of CO2 (Saxe et al. 1998).

Indeed, when both CO2 and water supply were manipulated in 11 epiphytic bromeliad species in a growth chamber study (Zotz et al. 2010), growth was only

stimulated by CO2 in a single species, while the majority of species showed the

expected effect of different water supply. With the exception of two species, the

expected mitigation of drought stress by elevated CO2 was not found.

To summarize, current evidence does not support the notion that elevated CO2

will substantially increase growth and water use efficiency among vascular

epiphytes. However, this conclusion is based on a very limited number of studies.

Considering that epiphytes have been called “especially vulnerable” to global

change (Benzing 1998), a much greater research effort seems warranted to allow

better predictions of the potential consequences of increased CO2 and other aspects

of global change for epiphytes. Last but not least, such experiments should also

address possible interactions of the abovementioned factors with increasing





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Box 5.4 Growth Analysis in Vriesea sanguinolenta (Stefan Wester, Cord Mikona, and Gerhard Zotz)

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