Tani et al.
Progress in Earth and Planetary Science(2024) 11:42
https://doi.org/10.1186/s40645-024-00645-8

Progress in Earth andPlanetary Science

Open Access

REVIEW

Basal emission rates of isoprene
and monoterpenes from major tree species
in Japan: interspecies and intraspecies
variabilities
Akira Tani1*, Noboru Masui1, Ting‑Wei Chang1, Motonori Okumura2 and Yutaka Kokubu3

Abstract
Uncontrolled terpenoid emissions from forest trees in Japan may have contributed to high O­ 3 concentrations
observed in urban and suburban areas. To estimate ozone formation via a series of reactions between ­NOx and ter‑penoids using atmospheric chemistry models, it is important to produce terpenoid emission inventories by collect‑
ing all reported emission data for the major tree species in Japan and examining their reliability. In this review, we
first describe three different plant terpenoid emission types, i.e., isoprene-emitting type, monoterpene-emitting
type with storage tissues and organs, and monoterpene-emitting type without storage tissues and organs. Second,
we describe various methods for measuring plant terpenoid emissions, including a recently developed simplified
method, and explain their reliability. We emphasized that applicable measurement methods depend on the terpe‑
noid emission types. Data obtained using static chamber methods should not be considered because they havethe highest uncertainty resulting from normal chamber materials that are not specific to terpenoid measurements
and lack humidity control. Finally, we show the absolute values of the collected emission rates and describe theirvariability. The deciduous oak species, Quercus serrata and Quercus mongolica var. crispula, and bamboo species, Phyllostachys pubescens and Phyllostachys bambusoides, are strong isoprene emitters. Among the monoterpene emit‑
ters, four evergreen broadleaf trees, including three Quercus species, had the highest basal emission rate (BER). The
monoterpene storage type conifers Larix kaempferi and Pinus densiflora have relatively lower BERs. Emission data are
not available for Castanopsis cuspidata, and seasonal changes in emission rates have not been reported for several
major tree species in the top 20 rankings. Within species, the reported emission rates of some tree species differedby threefold. These differences may be attributed to the reliability of the measurement and analytical systems, tree
age, leaf morphology, environmental conditions, and genetic diversity. We emphasize the need for reliable measure‑ments to achieve a more precise terpenoid emission inventory for major tree species in Japan.
Keywords Tower flux measurement, Flow-through chamber, Static chamber, Temperature, Light intensity, Quercus,
Bamboo*Correspondence:
Akira Tani
atani@u-shizuoka-ken.ac.jp
Full list of author information is available at the end of the article© The Author(s) 2024. Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which
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regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of thislicence, visit http://​creat​iveco​mmons.​org/​licen​ses/​by/4.​0/.

Tani et al. Progress in Earth and Planetary Science(2024) 11:42

1 Introduction
Higher plants produce terpenoids, including isoprene
and monoterpenes, as secondary metabolites. They are
major biogenic volatile organic compounds (BVOCs),
and their total annual emissions are estimated to be 600–
800 Tg C (Arneth et al. 2008). This value is higher thanthe estimated annual emissions of anthropogenic volatileorganic compounds (AVOCs) (98–158 TgC) (Boucher
et al. 2013). Given their high reactivity in the atmosphere(Atkinson and Arey 2003), terpenoids play a significant
role in the production of photochemical oxidants, including ozone (­O3), and secondary organic aerosols (SOA).
Tropospheric ­O3 negatively affects humans, animals, andplants (Tani and Mochizuki 2021; Masui et al. 2023). The
scattering and absorption of sunlight by SOA can influence global radiation, and cloud condensation nuclei
derived from SOA may also affect climate. Nevertheless,accurately quantifying the extent of their contribution
remains challenging, with significant uncertainties persisting (Shrivastava et al. 2017).
In the urban areas of Japan, the O­ 3 concentration
remains high despite the reduction in ­NOx and AVOC
emissions under the national air pollution control law. It
has been suspected that the uncontrolled BVOC emissions might contribute to the ­O3 formation (Churkina
et al. 2017; Gu et al. 2021). Thus far, BVOC emission models have been developed (Evans et al. 1985; Tingey et al.
1991; Guenther et al. 1993, 2006; Niinemets and Reichstein 2002, 2003) to explain VOC emission processes inplants. G93 model (Guenther et al. 1993) and Model ofEmissions of Gases and Aerosols from Nature (MEGAN)(Guenther et al. 2006) are widely used to describe leaflevel and large-scale VOC emissions, respectively. The
G93 model describes temperature-dependent monoterpene and temperature- and light-dependent isopreneemissions. MEGAN employs these functions as a basic
theory but includes the effects of cumulative temperatureand other environmental stresses. MEGAN can produce
a BVOC emission map and provide data for atmosphericchemical reaction models to estimate ­O3 formation.
There are considerable uncertainties in the BVOC
emission estimates owing to uncertainties in the data
input to G93 and MEGAN. The basic inputs are (1) biomass distribution (the fraction map of land cover by vegetation and the map of leaf area index [LAI]), (2) emission
rates (leaf area-based emission rate in G93 and groundarea-based emission rate in MEGAN), and (3) environmental factors, including light intensity, temperature, andenvironmental stresses. Among these inputs, (1) and (2)are the major factors affecting BVOC emission estimates(Guenther et al. 2006, 2012).
As the terpenoid emission rates are affected by temperature and light intensity, they are standardized

Page 2 of 18

to a photosynthetic photon flux density (PPFD) of
1000 μmol ­m−2 ­s−1 and a leaf temperature of 30 °C (i.e.,
BER). The BERs data for different tree groups or tree species are used as inputs for MEGAN and other emission
models. The values are not constant throughout the year
for both evergreen and deciduous trees and usually thehighest in summer, lower in spring and autumn, and thelowest in winter. However, exceptions have been reported
for some tree species (Tani et al. 2002; Matsunaga et al.
2013; Okumura et al. 2013) in Japan and the seasonality of terpenoid emission rates may differ for different
tree types. In addition, the reported BERs were obtainedusing different measurement methods, indicating that
their degree of reliability is dependent on the advantages
and disadvantages of the methods.
Several studies have reported the influence of input
data uncertainty on BVOC emission estimates (Kim et al.
2014; Wang et al. 2018; Batista et al. 2019). For example,
the global estimates of BVOC emission with MEGAN
can differ by − 9% and + 75% for isoprene and α-pinene,
respectively, when using updated basal emission rates(BERs) instead of the default MEGAN’s BERs (Henrot
et al. 2017). In China, using different vegetation land
cover data as inputs to MEGAN can result in a 52% difference in ­O3 concentrations (Ma et al. 2023).
In Japan, Kannari et al. (2007) developed a nationwide
annual BVOC emission inventory for Japan, namely
the East Asian Air Pollutant Emissions Grid Database(EAGRID2000-Japan), based on a procedure similar tothat of the BVOC emission model called the Biogenic
Emissions Inventory System (BEIS) (Pierce and Waldruff
1991). Although the emission inventory was made at afine resolution (1-km spatial and 1-h temporal resolutions), the land cover data of vegetation as inputs werecategorized into only two large general groups of trees(broadleaf and coniferous trees) and six crops (rice,wheat, beans, forage/manure crops, industrial crops, andothers). Thus, these data are still too coarse to properly
represent vegetation characteristics specific to Japan.
Further improvements in emission inventories have been
made in various studies. Inoue et al. (2010) improved
the EAGRID2000-Japan and estimated the nationwide
BVOC emission as 3312 Gg ­yr−1 based on the G93 algorithm by incorporating species-specific BERs of the
major vegetation (10 species) in the Kinki region of Japanderived from Bao et al. (2008). However, there exists
high uncertainty in the deduced emission value becausesome of the BERs (e.g., rice paddy) provided by Bao et al.
(2008) are suggested to be overestimated (Tani et al.
2023). A recent study of nationwide BVOC emissions(Chatani et al. 2015) demonstrated a lower emission
value of 2378 Gg y­ r−1 (isoprene 35%, monoterpenes 20%,
sesquiterpenes 10%, and other VOCs 35%) via MEGAN

Tani et al. Progress in Earth and Planetary Science(2024) 11:42

using species-specific BERs obtained from the latest literature. Chatani et al. (2018) also used updated data ofvegetation land cover and species-specific BERs (16 species) for MEGAN and obtained in a slight improvementin the model’s performance of air quality simulation forpredicting ­O3 concentrations. They used some of available BERs data including overestimated ones (Chatani
et al. 2015, 2018).
To produce a reliable inventory of tree terpenoid emissions, it is important to collect as much BERs from theliterature as possible and examine the reliability of thereported individual terpenoid emission rates obtainedusing a wide variety of methods. In this review, we
first describe different types of plant terpenoid emissions because the reliability of the measurement methods depends on the emission type. Second, we describe
various methods, including one recently developed, formeasuring plant terpenoid emissions and explain degreesof their reliability. Finally, we show the variabilities of thecollected emission rates, which are affected by tree origin
and age, growing conditions, and measurement methods, and emphasize the need for future measurements
to achieve a better terpenoid inventory of major tree
species. Crops in Japan might be terpenoid sources, andtherefore, terpenoid emission data from crops are alsocollected from the literature and shown in this review.

2 Terpenoid emission characteristics of plants
Plants synthesize terpenoids via secondary metabolism
and release them into the atmosphere. The major compounds emitted by plants are volatile terpenoids, including isoprene ­(C5H8) and monoterpenes ­(C10H16). Some
plants also emit sesquiterpenes (­C15H24) and diterpenes­(C20H32) which are semi-volatile. These compounds are
generally highly reactive and thus are highlighted by theirhigh ­O3-forming capacity in the atmosphere. Notably,
not all plants produce or emit these compounds. Here,
we briefly summarize these terpenoids and the commonly observed terpenoid-emitting plant species.
Isoprene (2-methyl-1,3-butadiene) is one of the mostremarkable compounds among the terpenoids emitted by plants, with a molecular weight of 68.12 and aboiling point of 34.07 °C. Typical isoprene-emitting
species are deciduous broadleaf trees such as Quercus
serrata, Quercus crispula, Populus spp., and bamboo(Stewart et al. 2003; Tani and Kawawata 2008; Tani
and Mochizuki 2021). The global annual emission isestimated to be 500 ± 100 × ­1012 g C y­ r−1 (Arneth et al.
2008). The isoprene precursor dimethylallyl diphosphate (DMAPP) is converted to isoprene by isoprenesynthase, which is embedded in the thylakoid membrane of the plastid (Sasaki et al. 2005). The rate ofisoprene emission typically increases with increasing

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temperature and light intensity, reaching a light saturation point of 200–1000 µmol ­m−2 ­s−1 depending onplant growth conditions (Fig. 1). Additionally, theoptimum temperature for isoprene synthase activity isapproximately 40 °C (Guenther et al. 1993).
Monoterpene is the general term of C
­ 10H16 terpenoid
compounds and sometimes includes oxygenated compounds such as monoterpene alcohols (e.g., linalool,­C10H18O) and monoterpene ketones (e.g., camphor,­C10H16O). Monoterpenes are the main components thatmake a plant “fragrant.” The typical isomer of monoterpenes emitted from plants is α-pinene, β-pinene, camphene, and limonene. The composition of monoterpene
isomers varies among plant species. Monoterpenes aresynthesized in the plastids of leaves and flowers. Geranyl diphosphate (GPP), a monoterpene precursor inplants, is synthesized from DMAPP and isopentenyl
pyrophosphate (Dudareva et al. 2005; Dong et al. 2016).
GPP is converted into monoterpenes by the synthases
corresponding to each monoterpene (Degenhardt et al.
2009). Plants that emit monoterpenes can be classified
into storage and non-storage types. The storage type
plants include most of the conifers such as Cryptomeria
japonica, Chamaecyparis obtusa, Pinus densiflora,
Larix kaempferi, and Picea jezoensis var. hondoensis.
The non-storage type plants are evergreen broadleaf
trees such as Quercus ilex, Quercus phillyraeoides,
and Castanopsis sieboldii var. sieboldii. Monoterpene
emissions from storage type plants typically follow atemperature-dependent evaporation process from theorgans in which monoterpenes are stored (e.g., resin
canals and oil glands) (Fig. 1) (Gershenzon et al. 2000).
In contrast, monoterpene emission in the non-storage
type plants depends on temperature and light intensity,
showing a pattern similar to that of isoprene emission
from isoprene-emitting plants. Notably, some plants
do not usually produce monoterpenes, but metabolic
pathways may be activated to produce them when stimulated by external stresses, such as feeding damage byinsects or mechanical damage.
During biosynthesis, farnesyl pyrophosphate and geranylgeranyl pyrophosphate are converted to sesquiterpenes and diterpenes, respectively (Davis and Croteau
2000). Their emission rates are generally influenced byabiotic factors such as temperature and light, as wellas biotic factors such as diurnal cycle, seasonality, andplant developmental stage (Duhl et al. 2008). Currently,
the emissions of sesquiterpenes and diterpenes fromplants are less well understood than those of isoprene
and monoterpenes because of difficulties in the sampling processes associated with the higher boiling point
and stronger adhesion of these semi-volatile compounds(Helmig et al. 2004; Duhl et al. 2008).

Tani et al. Progress in Earth and Planetary Science(2024) 11:42

Page 4 of 18

Fig. 1 Isoprene and monoterpene emission rates in response to environmental factors. Isoprene and monoterpene emission rates
from non-storage type plants are shown as relative to those at a temperature of 30 °C and a photosynthetically photon flux density (PPFD)
of 1,000 µmol ­m−2 ­s−1. Monoterpene emission rate from monoterpene storage type plants, which depends on temperature only in most cases,
is shown as relative to that at 30 °C. The figure was drawn using values calculated using so-called G93 algorism (Guenther et al. 1993)

3 Measurement methods for plant terpenoid
emissions
The methods used to measure the terpenoid emission
rates of trees are listed in Table 1. These methods canbe broadly classified into two types: tower flux andchamber methods. Tower flux method includes eddy
covariance (EC), relaxed eddy accumulation (REA),
and gradient flux (GF) methods. The chamber method
is classified into flow-through and static chamber
methods.
Tower flux method can provide large-scale observations of regional terpenoid emission characteristics. Thismethod is advantageous as it does not require physicalcontact with plant leaves and branches, and can thus
avoid mechanical stimuli that may induce the emission ofcertain species of monoterpenes, including linalool (Taniet al. 2023). The chamber method is used for the emission
measurement of potted trees, branches, and single leaves,and is easy to install on site. In this section, we describe
the characteristics, advantages, disadvantages, and reliabilities of these methods, because they may cause variability in the terpenoid emission rates of identical tree
species.

3.1 EC method

To conduct an EC measurement on a flux tower, thereal-time terpenoid concentration must be measured
above the forest canopy, coupled with the three-dimensional wind direction and velocity. High-time-resolution measurement instruments are required to measure
the concentrations. Proton transfer reaction mass spectrometry (PTR-MS) was used in many cases to obtaina concentration signal at approximately 10 Hz (Müller et al. 2010; Mäki et al. 2019; Sarkar et al. 2020).
Frequent calibrations must be conducted onsite using
standard gases. However, PTR-MS cannot precisely
discriminate the isomers of monoterpenes and produces fragment ions originating from them and other
VOCs that may interfere with accurate concentration
determination (Tani et al. 2003, 2004). In an atmosphere dominated by isoprene, m/z 69 is the ion mainly
produced from isoprene, facilitating more accurate
concentration determinations. Through the applicationof data processing techniques such as axis rotation, thismethod can be extended to encompass slopes and complex terrain.

LC method requires commercial port‑
able photosynthetic measurement
systems or man-made cuvette. Stomatal
conductance and net assimilation rate
can be obtained simultaneously. As
the commercial systems can control
light intensity and leaf temperature,basal emission rate can be directly
measured
FPC requires commercial environmen‑
tally controlled chambers or man-madeflow-through chambers

Plant canopy

Plant canopy

A branch with several leaves

A part of a leaf

Relaxed eddy accumulation (REA)

Gradient flux (GF)

Bag enclosure (BE)

Leaf cuvette (LC)

Flow-through plant chamber (FPC) Individual plant

The method cannot be applied to slopeand complex terrain. Axis rotation
is not efficient if horizontal wind direc‑
tion is not constant

A proton transfer reaction mass
spectrometry (PTR-MS) is used
to obtain the concentration signal.
Isomers of monoterpenes and frag‑ment ions originating from monoter‑penes and other VOCs may interferewith the accuracy of the determined
concentration

Key points and constraints

A branch is enclosed with a transparent
bag made from fluorine-based resin. The
lid of the bag is tightened, and clean air
is sent to the sealed bag

LAI is required

Coefficient (leaf area index [LAI] or specificleaf weight [SLW]) to convert the rate
on leaf area or dry weight basis to that onland area basis is required

Emission data on land area basis are
obtained. They can be used as inventory
data without unit conversion

Emission data on land area basis are
obtained. They can be used as inventory
data without unit conversion

Emission data on land area basis are
obtained. They can be used as inventory
data without unit conversion

Conversion to emission inventory data

Flow rate must be adjusted to avoid
Coefficient (SLW or LAI) to convert the ratewater condensation inside. Terpe‑on leaf area or dry weight basis to that onnoid adsorption on the inner surface
land area basis is requiredof the chamber may not be negligible
if the chamber is not made using propermaterials. As a young sapling is usedin most cases, scaling up of the emission
rate to that of an adult tree should be
carefully performed

Replicate measurements are required
to obtain representative basal emis‑
sion rate. Mechanical stimulus may
induce temporary terpenoid emissions
from monoterpene storage type trees

Water vapor should be removed
in the air preparation process. Touching
stimulus may cause unexpected VOCemissions

Terpenoid concentrations are measured This method is valid only under neutral
at two heights above the forest canopy. and unstable atmospheric conditions
Flux is calculated from the concentration
gradient and eddy diffusion coefficient

Real-time sampling is required depend‑ing on vertical wind direction

Measurement instruments with hightime resolution (~ 10 Hz) are required
to measure the concentration. The
method can be applied to slopeand complex terrain

Plant canopy

Eddy covariance (EC)Measurement technique

Measurement object

Measurement method

Table 1 Measurement methods for terpenoid emission rates and their characteristics

Tani et al. Progress in Earth and Planetary Science(2024) 11:42
Page 5 of 18

Excised leaf tip (~ 1 × 2 cm)

Leaf-tip vial (LV)

A leaf tip in a vial is incubated at con‑stant temperature and light intensity
for a certain period (30–60 min),followed by the sampling of the gas
inside. The LV method is suitable for fast
screening of isoprene emission capacity
as simultaneous measurements can be
conducted for multiple vials

A whole plant or a part of a plant The SC method uses commercial
or man-made closed chambers

Static chamber (SC)Measurement technique

Measurement object

Measurement method

Table 1 (continued)

Excising leaf tip may induce terpenoid
emission from monoterpene storage
organs. A cross check of this methodwith more reliable methods suchas the LC method is required

Water vapor originating from planttranspiration is condensed on the innersurface without removing water
vapor in the chamber. This resultsin the loss of water-soluble VOCs. As
most of the chambers are not madeusing special materials, adsorption
and desorption of terpenoids may
occur on the surface. As a young
sapling is used in most cases, scaling-upof the emission rate to that of an adulttree should be carefully performed

Key points and constraints

Coefficient (SLW or LAI) to convert the rate
on leaf area or dry weight basis to that onland area basis is required

Coefficient (SLW or LAI) to convert the rate
on leaf area or dry weight basis to that onland area basis is required

Conversion to emission inventory data

Tani et al. Progress in Earth and Planetary Science(2024) 11:42
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Tani et al. Progress in Earth and Planetary Science(2024) 11:42

3.2 REA method

Similar to the EC method, the REA method measures theregional flux on a tower, but it does not require real-timeor high-time-resolution concentration measurements.
The upward and downward air is separately sampled into
adsorbent tubes or evacuated canisters according to thevertical wind direction (Mochizuki et al. 2014; Sarkar
et al. 2020). Because real-time sampling according to thevertical wind direction is required, this method may not
be applicable to slopes and complex terrains unless thehorizontal wind direction is almost constant, which is
not realistic in the field.
3.3 GF method

The GF method is a basic method for determining thefluxes of trace gases, including terpenoids. Terpenoid
concentrations are measured at two heights above theforest canopy, and the flux is calculated from the concentration gradient and eddy diffusion coefficient (Taniet al. 2002; Schween et al. 1997). The eddy diffusion coefficient can be determined using the vertical profile ofwind velocity or heat balance above forests. This method
is only valid under neutral and unstable atmosphericconditions.
3.4 Bag enclosure method

The bag enclosure (BE) method is a flow-through chamber method (Guenther et al. 1993; Owen and Hewitt
2000; Okumura et al. 2008b). A branch with several
leaves is enclosed in a transparent bag made of fluorinebased resin. The lid of the bag is tightened, and clean air
is sent to the sealed bag at a flow rate of 5–10 L m­ in−1
depending on the total leaf area. Clean air is produced
using activated charcoal or a heated platinum catalyst. It
is better to remove water vapor during the air preparation
process to avoid water condensation on the inner surfaceof the bag caused by the water vapor increment by planttranspiration. This prevents VOC dissolution into condensed water. Physical contact with branches and leaves
can trigger VOC emissions. Therefore, it is important toallow the branch to rest for a designated period ranging
from several hours to a day to mitigate such emissions.
Because the emission rate is determined based on leaf
area or dry weight, a coefficient is necessary to convert
this rate to that on a land area. This coefficient represents
LAI or the specific leaf weight.
3.5 Leaf cuvette method

The leaf cuvette (LC) method is a flow-through chamber method that uses commercial portable photosynthetic measurement systems (Pétron et al. 2001; Tani
and Kawawata 2008) or hand-made cuvettes (Monson
and Fall 1989; Lehning et al. 1999). A portion of the air

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outflowing from the leaf cuvette is collected into adsorbent tubes. Some of the latest systems already have anembedded sampling port (e.g., LI-6800, Li-Cor Inc., Lincoln, NE, USA), whereas modifications are needed forolder systems (e.g., LI-6400, Li-Cor Inc.). As these systems can control environmental factors, such as lightintensity and leaf temperature, and measure stomatal
conductance and net assimilation rate, it is possible toinvestigate the effects of environmental factors on terpenoid emission rates and address the relationships
between plant physiological parameters and emission
rates. It is easy to directly obtain BER, defined as theemission rate normalized to 30 °C leaf temperature and1000 µmol ­m−2 ­s−1 PPFD (Guenther et al. 1993), by controlling the measurement conditions. However, chamber wall, gasket surfaces, and O-rings may potentiallycause adsorption/desorption problems for VOC studies(Niinemets et al. 2011).
Obtaining representative BERs of sunlit and sun shaded
leaves necessitates conducting replicate measurements ofindividual trees. As the measured leaf is held down by agasket to seal the cuvette, this mechanical stimulus may
induce temporary terpenoid emissions from monoterpene storage type trees. LAI is required to convert theemission rate from a leaf area basis to a land area basis.
3.6 Flow‑through plant chamber method

The flow-through plant chamber (FPC) method uses
commercial environmentally controlled chambers or
man-made flow-through chambers (Kim et al. 2001; Tani
et al. 2023) for the entire plant. In this method, water
condensation can be avoided by controlling humidity.
However, with an increase in the number of plant leaves
higher rates of air inflow into the chamber are required
to avoid water condensation. If the chamber is not madeof special materials to avoid VOC adsorption, terpenoid
adsorption on its inner surface may not be negligible,
particularly for oxygenated monoterpenes and less-volatile terpenoids. In this case, desorption of previously
adsorbed terpenoids from the chamber may also occur,
resulting in an increased uncertainty in the obtained
emission rates. As potted plants are used in many cases,
the emission rate on a plant basis needs to be converted
to that on a leaf area basis or dry weight basis, and thereafter, the value is recalculated on a land area basis. When
young saplings are used for measurement, the emission
rate should be carefully scaled up to one of the adult trees
because the emission rate may be different for trees ofdifferent ages (Kim et al. 2005).
3.7 Static chamber method

The static chamber (SC) method uses commercial or
hand-made closed chambers for the entire plant and part

Tani et al. Progress in Earth and Planetary Science(2024) 11:42

of a plant (Kim et al. 2005; Bao et al. 2008; Brennan et al.
2022). Without removing water vapor from the chamber,water vapor originating from plant transpiration condenses on the inner surface of the chamber, resulting in
the loss of water-soluble VOCs. Even if heat exchangers
are used to remove water vapor, VOCs dissolve into liquid water condensed on the cooled fin. In some commercially available closed chambers, environmental factors,
such as light intensity, air temperature, and humidity, canbe controlled (Bao et al. 2008). As most of the chambers
are not made of special materials to avoid VOC adsorption, attention should be paid to the adsorption and desorption of terpenoids. As young plants are used in manycases, the scaling up of the emission rate follows the samemanner as described in the FPC method. The unit conversion follows that of the FPC method. The SC method
has the highest uncertainty, and therefore, it is not preferred for determining terpenoid emission rates.
3.8 Leaf‑tip vial method

The leaf-tip vial (LV) method uses an excised leaf tip(~ 1 × 2 cm) to determine the terpenoid emission rate
(Chang et al. 2022b). The leaf tip is placed horizontally ina laid-down vial (10 mL volume) and incubated at a constant temperature and light intensity for a certain period(30–60 min), followed by the sampling of the gas inside.
The LV method is particularly suitable for fast screening of isoprene emission capacity because measurements
can be conducted almost simultaneously for multiple
vials (5–10 vials). To validate the LV method for a target plant, it must be cross-checked with more reliable
methods such as the LC method. If a good correlation,such as a linear regression, is observed for the terpenoid
emission rates determined by the LV and LC methods,
the LV method is judged to be applicable. Mechanicalwounding by cutting a leaf tip may induce monoterpene
emissions from monoterpene storage organs; therefore,
the LV method is not suitable for monoterpene-storage
plants. Currently, this method is applicable to the measurements of isoprene emission from palm trees (Changet al. 2022b) and monoterpene emissions from Q. phillyraeoides (Chang et al. 2024).

4 Terpenoid emission data of trees and crops
grown in Japan
Data on BERs were collected from published literatureand our original experiments (unpublished). The target
tree species were selected from the top 20 dominant tree
species in Japan, based on a survey by the Japanese Forestry Agency (Table 2). Terpenoid emission data of major
crop species are also added. The BERs are defined as theemission rate normalized to 30 °C leaf temperature and1000 µmol ­m−2 ­s−1 PPFD for both isoprene emitters andPage 8 of 18

non-storage types of monoterpene emitters and to 30 °C
leaf temperature without any specific light conditions forthe storage type monoterpene emitters (Guenther et al.
1993).
4.1 Deciduous broadleaf trees
4.1.1 Monoterpene emitters

Many wild Acer species occur in Japan, and many ofthem are known to be monoterpene emitters. Acer species are categorized as a group in the dominant tree ranking in Japan and ranked 11th. Mochizuki et al. (2020a)
reported monoterpene BERs of Acer and Fagaceae tree
species grown in Japan. Six Acer tree species (A. palmatum, A. japonicum, A. sieboldianum, A. shirasawanum,
A. rufinerve, and A. micranthum) were identified as
monoterpene emitters (Table S1). Seasonal variation in
the monoterpene emission rate of A. palmatum, a nonstorage type monoterpene emitter, was measured using
the LC method (Mochizuki et al. 2020b; Figure S1),
which showed that it increased from April, peaked inAugust, and decreased in winter. The dominant component was α-pinene followed by sabinene and β-pinene. In
other studies, A. platanoides, A. campestre, and A. buergerianum have also been shown as monoterpene emitters(Acton et al. 2016; Baraldi et al. 2019; Chang et al. 2021a;
Table S1). Although the seasonal trend of monoterpene
emissions from A. palmatum may offer insights applicable to other Acer species, it is advisable to gather additional data on various Acer species. Currently, seasonalvariation data on monoterpene BERs are limited to A.
palmatum.
In contrast, monoterpene emissions were not detected
in other 14 Acer species (A. amoenum, A. tenuifolium, A.
crataegifolium, A. pictum, A. cissifolium, A. argutum, A.
distylum, A. nipponicum, A. miyabei, A. maximowiczianum, A. tataricum, A. pycnanthum, A. diabolicum, andA. carpinifolium) and two Fagus species (F. crenata andF. japonica) (Table S1), even in summer. However, A. pictum (19th rank in in the dominant tree species) occurring in China (Klinger et al. 2002) and Korea (Chang et al.
2021a) has been reported to emit monoterpenes, indicating a discrepancy in terpenoid emission trends between
Japan and other Asian countries.
Castanea crenata, 14th rank in the dominant tree species, was reported as a monoterpene (e.g., α-pinene,
sabinene, and β-pinene) emitter with a BER of
1.1 nmol ­m−2 ­s−1 obtained using the LC method (Mochizuki et al. 2020a). However, the peak and seasonal variations in the monoterpene BERs of C. crenata have notbeen reported.
Betula trees in Japan are reported to be monoterpene
emitters (Yang et al. 2021; Masui et al. 2022). For example, B. platyphylla var. japonica has a monoterpene BER

Tani et al. Progress in Earth and Planetary Science(2024) 11:42

Page 9 of 18

Table 2 Top 20 ranking dominant tree species in Japanese forest and their terpenoid emissions characteristics
Tree species

Rank Total area ­(103 ha) Data availability of basal emission rate
Isoprene in summer

Monoterpene in summer

Value

Type** Value

Unit

Unit

Seasonalityβ

Cryptomeria japonica

14369.60S< 0.1–5.2 μg ­g−1 ­h−1

0.03–0.17

Chamaecyparis obtusa

22730.40S

0.1–1.0μg ­g−1 ­h−1

0.08–0.12

Quercus serrata

31596.020.5–42.9

nmol ­m−2 ­s−1

6.4–114.9μg ­g−1 ­h−1

●
●
●

0

Abies sachalinensis

41330.40

NA

NA

Other broad-leaved trees

51131.2***Pinus densiflora61092.80

Fagus crenata

7

1051.2[Non-emitterof terpe‑
noids]

Quercus mongolica var.
crispula

8

972.0

26.3–29.5

Larix kaempferi

9

876.0

0

Betula ermanii

10

640.0

0

S

0.6–2.5

nmol ­m−2 ­s−1 < 0.01–0.19 ●S

1.5–1.9μg ­g−1 ­h−1nmol ­m−2 ­s−1

0.12–0.14●

0S

1.2–1.7

nmol ­m−2 ­s−1 0.09 -0.20

NA

●
NA

Acer spp.

11

313.6

0

Tilia japonica

12

277.6

0

S

1.0

*
μg ­g−1 ­h−1*

Castanopsis cuspidata var.
sieboldii

13

273.6

0

NS

15.0

nmol ­m−2 ­s−1

Castanea crenata

14

239.2

0

1.1

nmol ­m−2 ­s−1

Alnus spp.

15

227.2

0

*

0.10

NA
NA
NA
NA

Carpinus spp.

16

200.8

*

Quercus glauca

17

192.0

0

*

Castanopsis cuspidata

18

184.8

NA

Acer pictum

19

182.4

< 0.1

μg ­g−1 ­h−1

Phyllostachys pubescens

20

181.6

12.8–22.8

nmol ­m−2 ­s−10●Phyllostachys bambusoides

24

115.2

17.6

nmol ­m−2 ­s−1

0

●

NS

8.0

*
nmol ­m−2 ­s−1

NA
NS

0–3.3

NA
NA

μg ­g−1 ­h−1

NAThis table is arranged based on a dataset of "Forestry Agency Forest Ecosystem Diversity Basic Survey, Phase 3 (2009–2013 at 13,357 survey sites).” The survey sites
were divided into a 4-km square grid, and the intersection of the squares with 0.1 ha area was surveyed for the dominant tree species. The dominant tree species hasscore of “1” (one dominant tree species: 11,719 sites) or “0.5″ (two dominant tree species: 1,631 sites)
The total area of trees was calculated by multiplying the accumulated score of each tree species by 16 k­ m2, assuming that a 4 km square around a survey site has thesame tree composition
The BER of isoprene type and monoterpene non-storage type is a normalized rate under the standard condition of 1,000 µmol m­ −2 ­s−1 PPFD and 30 °C leaf
temperature. The BER of monoterpene storage type is a normalized rate under the standard condition of 30 °C leaf temperature. For monoterpene storage type,temperature dependence coefficient β was experimentally determined in most cases●: Seasonality data are available, NA: not available, 0: no emission. *The availability of emission data depends on the tree species. **S: monoterpene storage type, NS:monoterpene non-storage typeThe detailed information including references, measurement techniques, and tree age for each data is provided in supporting information S1–S3 and Table 3

of 2.8 nmol ­m−2 ­s−1 in July (β = 0.09; Masui unpublished)
as determined by the BE method. B. ermanii is the 10th
ranked dominant tree species in Japan; however, dataon whether it is a monoterpene emitter are unavailable.
However, the impact of BVOC emissions on ­O3 formation in urban areas seems to be negligible because Betulaspp., including B. ermanii, are found only in the northern and alpine regions. Tilia japonica is ranked 12th in
the dominant tree species and is found to be a monoterpene emitter with a BER of 0.97 µg ­g−1 ­h−1 (Jing et al.

2020); however, it also occurs in the northern and alpine
regions.
4.1.2 Isoprene emitters

Data on isoprene BERs of Quercus species are available
in the literature (Bao et al. 2008; Okumura et al. 2008c;
Tani and Kawawata 2008; Lim et al. 2011; Kim and Lee
2012; Chang et al. 2021a) and are summarized in Table 3.
Some studies have reported seasonal variations in isoprene emission rates of Q. mongolica var. crispula (Kim

Tani et al. Progress in Earth and Planetary Science(2024) 11:42

Page 10 of 18

Table 3 Basal emission rates (BERs) of isoprene reported for deciduous Quercus trees
Tree species

Age

MethodSeasonBER

Citation

Quercus mongolica var. crispula(Adult tree)

BE

Jun.
Jul.
Aug.
Sep.
Oct.

4.70
29.50
26.30 nmol m 2 s 1
10.90
2.90

Matsunaga et al. (2012)

3 yearsLC

May–Jul.

27.30 nmol m­ −2 ­s−1

Tani and Kawawata (2008)

3–5 years

SC–

26.0 ug ­g−1 h−1

Bao et al. (2008)

3 yearsLC

May–Jul.

27.80 nmol m­ −2 ­s−1

Tani and Kawawata (2008)

40 years

LC (sunlit leaf )LC(sunshade leaf )

Jun.

42.90
20.50 nmol ­m-2 ­s-1

Okumura et al. (2008c)

3 years

FPC–

6.39 ug ­g−1 h−1Chang et al. (2021a)

3–5 years

SC–

224.2 ug ­g−1h−1

Bao et al. (2008)(10m height)

FPC

Spring64.0 ug g­ −1 h−1

Kim and Lee (2012)

SummerFall

79.5 ug ­g−1 h−1
60.1 ug ­g−1 h−1

Spring147.90 ug g
­ −1h−1

Quercus serrata

Lim et al. (2011)21–50 years

BE

SummerFall

114.92 ug g
­ −1h−1
114.92 ug g
­ −1h−1

3 yearsLC

May–Oct.

18.30 nmol m­ −2 ­s−1

Tani and Kawawata (2008)(middle age)BE

Jul.–Sep.

2.46 ug ­g−1 h−1Jing et al. (2020)

21–50 years

BE

May–Jun.

72.31

Lim et al. (2011)

BE

Aug.

89.31

BE

Oct.

11.14 ug ­g−1h−1

21–30 years

BE BE

May–Aug.

88.40
74.80

Quercus dentata

3 yearsLC

May–Oct.

30.00 nmol m­ −2 ­s−1

Tani and Kawawata (2008)

Quercus acutissima(10m height)BE

Spring1.35 µg ­g-1h-1

Kim and Lee (2012)

Summer Fall

1.71 ug ­g−1h−1
0.88

Quercus aliena

Quercus variabilis

Quercus mongolica

3 yearsLC

May–Oct.

0.00 nmol m­ −2 ­s−1

Tani and Kawawata (2008)

3 years

FPC

Aug.–Sep.

0.02 ug ­g−1h−1

Chang et al. (2021a)

3–5 years

SC–

0.18 ug ­g−1h−1

Bao et al. (2008)

41–50 years

BE

Jun.–Jul.

0.04 µg ­g-1h-1

Lim et al. (2011)

Sep. Oct

0.07 ug ­g−1h−1
0.05

3 yearsLC

May-Oct.

0.00 nmol ­m−2 ­s−1

3 years

FPC–

0.00 ug ­g−1h−1

Chang et al. (2021a)

15–20 years

BE

SpringSummerFall
Winter2.14
5.63
2.19 μg ­g-1h-1
0.00

Chen et al. 2020

11–50 years

BE

May–Jun.

72.99 µg ­g-1h-1

Lim et al. (2011)

BE

Aug.

65.85 µg ­g-1h-1

BE

Oct.

8.00 ug ­g−1h−1

BE
BE

May–Aug.

78.20
62.33

BE

Spring13.6 ug ­g−1 ­h−1

SummerFall

15.2 ug ­g−1h−112.0 ug ­g−1 h−1

21–30 years 41–50 years(10 m height)

Tani and Kawawata (2008)

Kim and Lee (2012)

The BER of isoprene type and monoterpene non-storage type is a normalized rate under the standard condition of 1000 µmol m­ −2 ­s−1 PPFD and 30 °C leaf
temperature
NA not available, LC leaf cuvette method, BE branch enclosure method, FPC flow-through plant chamber method, SC static chamber method
Q. mongolica is not endemic to Japan, but its BER is shown here for comparison purpose

Tani et al. Progress in Earth and Planetary Science(2024) 11:42

and Lee 2012; Matsunaga et al. 2012) and Q. serrata(Lim et al. 2011; Kim and Lee 2012). In Q. mongolica
var. crispula, isoprene emission is the highest in summer and lower in spring and fall (Table 3). This trend has
also been observed in other Quercus species such as Q.
aliena (Lim et al. 2011) and Q. mongolica in South Korea(Lim et al. 2011; Kim and Lee 2012). The isoprene BER
of Q. serrata is highly variable even for the same tree age(6.39 µg ­g−1 ­h−1;Chang et al. 2021a) to 224.2 µg ­g−1 ­h−1
(Bao et al. 2008), and the average value is higher thanthat of Q. mongolica var. crispula. The isoprene BER on
a land area basis was determined using the REA method
to be 13.3 nmol ­m−2 ­s−1 for a needle-broad mixed forest
including P. densiflora, Q. serrata, and Q. mongolica var.
crispula (Mochizuki et al. 2020c). The isoprene was estimated to be emitted from Q. serrata (2.2% based on total
basal area) and Q. mongolica var. crispula (1.0%).
F. crenata, 7th in the dominant tree species, wasreported as a non-emitter of isoprene and monoterpenes(Mochizuki et al. 2020a). We obtained the same results
using the LV and LC methods (unpublished). An exception was reported by Bao et al. (2008), who employed the
SC method using a commercial chamber and obtained an
isoprene BER of 0.79 μg g­ −1 ­h−1.

Page 11 of 18

4.4 Evergreen coniferous trees

In most cases, monoterpene emissions from coniferous
trees depend on leaf temperature. Data on the monoterpene BERs of the dominant evergreen coniferous trees,
C. japonica (1st), Chamaecyparis obtusa (2nd), and P.
densiflora (6th) are shown in Table S3 described individually in the following subsections.
4.4.1 Pinus densiflora

The monoterpene flux of a forest of P. densiflora(with 95% dominance) was measured by Tani et al.
(2002) using the GF method (Table S4). Major components of the monoterpene emissions from P.
densiflora were α- and β-pinene, limonene, andβ-phellandrene. The monoterpene basal flux was calculated using a fixed β-value of 0.19. It increased from
May (1.32 nmol ­m−2 ­s−1) to June (3.76 nmol ­m−2 ­s−1),
decreased below 1.0 nmol ­m−2 ­s−1 in summer, andincreased again to 5.28 nmol ­m−2 ­s−1 in October andNovember. The monoterpene basal flux of the same tree
species (with 84.5% dominance) in another forest measured using the REA method in August was reported tobe 2.50 nmol ­m−2 ­s−1 (Mochizuki et al. 2020c), which
is not largely different from that reported by Tani et al.
(2002).

4.2 Evergreen broadleaf trees

Some evergreen Quercus and Castanopsis trees emit
monoterpenes (Okumura et al. 2008b; Mochizuki et al.
2020a; Table S2). They are non-storage type monoterpene emitters, and their emissions depend on light intensity and leaf temperature. Particularly, C. cuspidata var.
sieboldii (13th rank) and Q. glauca (17th rank) have highmonoterpene BERs of 15.0 and 8.0 nmol ­m−2 ­s−1, respectively (Mochizuki et al. 2020a). Data for the 18th ranked
C. cuspidata, a species related to C. cuspidata var. sieboldii, are not available.
4.3 Deciduous coniferous trees

In Japan, deciduous coniferous tree species are limited to
L. kaempferi and some exotic trees such as Ginkgo biloba
and Metasequoia glyptostroboides. Among these tree species, L. kaempferi is 9th in the dominant tree species andhas been reported to be a monoterpene emitter (Iedaet al. 2006; Mochizuki et al. 2014). The monoterpene
emission from L. kaempferi depends on the leaf temperature, and the most dominant compound is α-pinene followed by sabinene, β-pinene, myrcene, and camphene(Mochizuki et al. 2014). Using the REA method, Mochizuki et al. (2014) obtained an almost constant value(~ 1 nmol ­m−2 ­s−1) of averaged BER standardized to 30 °C
for an L. kaempferi plantation from the bud break period
to fall (Figure S2, Table S3).

4.4.2 Cryptomeria japonica

C. japonica has three natural populations, including
Japan Sea, Pacific Ocean, and southwestern. Matsunaga et al. (2013) reported that seasonal variations in themonoterpene BER of C. japonica measured using the BE
method differed at different sites (Figure S3). For example, C. japonica individuals in Tanashi (around 50 years
old), a Pacific Ocean side population, had a monoterpene BER of 0.3 µg ­g−1 ­h−1 through a whole year except
for summer in which it was < 0.1 µg ­g−1 ­h−1. In contrast, adecreased emission rate in summer was not observed forthe southwestern population of Shiiba. Okumura et al.
(2013) employed the BE method for the Japan Sea side
population and found the highest monoterpene BER in
fall without suppression in summer (Figure S3, Table S5).
Hiura et al. (2021) obtained the monoterpene BERs
of C. japonica collected from 12 different planting sites
in three populations. In this study, three saplings of C.
japonica were transplanted from each planting site to anursery in the Miyagi Prefecture (38.78° N, 140.73° E).
Three years later, BVOCs emitted from the saplings werecollected using the BE method. The BER was largely different among the planting sites and even in identical populations (Figure S4). This study and the above-mentioned
studies suggest that differences in the reported monoterpene BERs may be due to intraspecies variations.

Tani et al. Progress in Earth and Planetary Science(2024) 11:42

All data collected by Okumura et al. (2013) are plotted against the leaf temperature (Figure S5). A clear
temperature dependence of monoterpene BER on
leaf temperature (β = 0.09, BER = 0.65 µg ­g−1 ­h−1,
RMSE = 0.93 µg ­g−1 ­h−1; β = 0.10, BER = 0.74 µg ­g−1 ­h−1,
RMSE = 0.88 µg ­g−1 ­h−1) was observed, suggestingthat it can be approximately estimated using leaf temperature without considering seasonal variations andpopulations.
4.4.3 Chamaecyparis obtusa

Matsunaga et al. (2013) employed the BE method andobserved relatively constant values of the monoterpene
BER of C. obtusa from spring to summer but a higher
value in winter. There were no differences in the monoterpene BERs between the Tanashi and Shiiba sites fromspring to fall (Figure S6). However, Okumura et al. (2008a)found high variability in monoterpene BERs among trees
at the same growing site (Figure S6, Table S6). By plotting all data (Figure S7) collected by Okumura et al.
(2008a), clear temperature dependence of the monoterpene emissions of a C. obtusa individual (Tree B) was
observed throughout the year (Figure S7; β = 0.09,
BER = 0.15 µg ­g−1 ­h−1, RMSE = 0.46 µg ­g−1 ­h−1; β = 0.10,
BER = 0.17 µg ­g−1 ­h−1, RMSE = 0.48 µg ­g−1 ­h−1). However,the correlation between temperature and monoterpene
emission rate was poor in “Tree A” (Figure S7; β = 0.09,
BER = 0.63 µg ­g−1 ­h−1, RMSE = 1.35 µg ­g−1 ­h−1; β = 0.10,
BER = 0.72 µg ­g−1 ­h−1, RMSE = 1.31 µg ­g−1 ­h−1). It may
be plausible to use the median or arithmetic mean of the
BERs; however, a relatively high error should be considered because of the high variation in the monoterpene
BERs among the individual trees.
4.5 Bamboo

In Japan, bamboo forests accounted for 0.6% of the total
forest area in 2012 (Forest Agency, 2018), and this area
has been increasing (Suzuki 2015). Most native or naturalized bamboo species in Japan belong to the Arundinarieae family. Only Kyushu Island and the Okinawa Islands
have large populations of Bambuseae species. Among thespecies of Arundinarieae, Phyllostachys pubescens, Phyllostachys bambusoides, and Phyllostachys nigra var. henonis are dominant bamboo species in Japan, which are
ranked as 20th, 24th, and 58th among the dominant tree
species, respectively.
Chang et al. (2021b) demonstrated that the isoprene BER of P. pubescens had a maximum of
22.8 ± 7.0 nmol ­m−2 ­s−1 in August. It then quickly
decreased to approximately one-tenth of its initial value
in autumn and completely seized by January. The emission increased back to a rate of 2.2 ± 0.2 nmol ­m−2 ­s−1
in May. Additionally, Chang et al. (2022a) measured

Page 12 of 18

isoprene BER of below-canopy culms in a pure P. pubescens forest using the LC method and determined the
BER to be 12.8 ± 5.7 nmol ­m−2 ­s−1 in August, with astrong correlation with specific leaf weight. Observation
data of tower flux measurements for P. pubescens vegetation are currently absent in Japan, but in China, Bai et al.
(2016) employed the REA method above a P. pubescens
forest and revealed a regional-scale isoprene BER of
0.42 mg ­m−2 ­h−1 (1.71 nmol ­m−2 ­s−1) in autumn (November 2012).
For P. bambusoides, Chang et al. (2021b) determined
the isoprene BER to be 17.6 ± 1.3 nmol ­m−2 ­s−1 in August.
For P. nigra var henonis, Chang et al. (2021b) obtained theisoprene BER of 5.0 ± 3.9 nmol ­m−2 ­s−1 in August.
4.6 Understory vegetation

In addition to canopy trees, understory vegetation, suchas herbaceous species and shrubs, should be considered when estimating BVOC fluxes in forest ecosystems.
Usually, the light intensity in the understory layer andforest floor is lower than that in the upper layer, which
may cause decreased BVOC emission rates. However,even in such environments, BVOC emissions from theunderstory vegetation may not be negligible in somespecific cases. Isoprene flux (> 0.5 nmol ­m−2 ­s−1) above
a L. kaempferi plantation in summer originated froman understory fern species Dryopteris crassirhizoma(Mochizuki et al. 2014).
4.7 Crops
4.7.1 Rice

The rice cultivation area in Japan is 1,404,000 ha (ThePortal Site of Official Statistics of Japan, 2021), which lies
between the 3rd and 4th dominant tree-growing areas(Table 2). Several studies have been conducted to qualitatively and quantitatively assess terpenoid emissions
from rice paddies. Winer et al. (1992) employed the BE
method in a rice paddy in California and found no emission of isoprene or monoterpenes from rice cultivar“M202.” Redeker et al. (2003) employed the SC method
for rice cultivar “Cocodrie” in a rice paddy and detected
no emission of monoterpenes, but detected a trace
quantity of isoprene. The isoprene emission rate converted on a land area basis was very low (3.7 µg ­m−2 ­h−1
≈ 0.015 nmol ­m−2 ­s−1). Tani et al. (1999) used the FPC
method for four cultivars grown in pots and reported
that a trace quantity of linalool was emitted from a ricecultivar “Mutsuhomare,” but not from the other threecultivars. Recently, Tani et al. (2023) observed linalool
emission from three Japanese rice cultivars using the
BE method. They attributed this emission to touching
stimuli by hand while enclosing the aboveground partsof the rice in the enclosure bag. Thereafter, Tani et al.

Tani et al. Progress in Earth and Planetary Science(2024) 11:42(2023) employed the REA method to measure the terpenoid flux above a rice paddy and reported a low quantityof α-pinene emission. The obtained flux of α-pinene was
0.006 nmol ­m−2 ­s−1 on a land area basis, with a 95% confidence interval of 0.004 nmol ­m−2 ­s−1.
By contrast, Bao et al. (2008) employed the SC method
using a commercial chamber for Japonica rice (cultivar
information was not provided). They found emissionsof five species of monoterpene (α-pinene, limonene,
myrcene, p-cymene, and β-pinene). Tani et al. (2023)
converted this emission rate on dry weight basis to thaton land area basis and reported that it corresponded to
600 µg ­m−2 ­h−1 (1.2 nmol ­m−2 ­s−1).
Except for Bao (2008), the other four studies reported
very small or negligible terpenoid emissions from rice
plants. As the REA method can avoid mechanical stimuli and is more reliable compared to other methods thatmight induce monoterpene emissions by mechanicalstimuli, the terpenoid emission rate seems to be very low.
4.7.2 Other crops

Wheat is the 2nd most dominant crop in Japan, with acultivation area of 220,000 ha (The Portal Site of Official Statistics of Japan, 2021), lying between the 15th and16th most dominant tree areas (Table 2). Some studies
conducted abroad provided the BERs of isoprene andmonoterpenes at 0–0.16 and 0–0.006 µg ­g−1 ­h−1, respectively (Evans et al. 1982; Winer et al. 1992; König et al.
1995; Morrison et al. 2016; Gonzaga Gomez et al. 2019).
Soybean is the 3rd most dominant crop in Japan, with acultivation area of 146,200 ha (The Portal Site of Official
Statistics of Japan 2021), lying between 21st and 22nd
dominant tree areas (Table 2). Isoprene and monoterpene emissions from soybean are negligible (Li and Sharkey 2013). Buckwheat is the 4th most dominant crop,
but its cultivation area (65,500 ha) is lower than that of
the 40th tree species and no report on its isoprene ormonoterpene emission is available.
Potatoes (70,900 ha), cabbage (34,300 ha), sweet potatoes (32,400 ha), and radish (29,200 ha) are widely cultivated (The Portal Site of Official Statistics of Japan2021). Cabbage is reported to emit monoterpenes at a
rate of < 1 µg ­g−1 ­h−1 (Vuorinen et al. 2004). Reports on
terpenoid emissions from the other three crops are notavailable.
4.8 Others

Chrysanthemum is the most widely cultivated cut flower(425,800 ha) followed by lily (65,900 ha). However, there
are no reports on terpenoid emissions from their leaves.
Pasture plants are grown in rural areas (717,600 ha).
Oxygenated alcohols have been reported to be emitted
from pasture plants, but the emissions of isoprene andPage 13 of 18

monoterpenes are negligible (Kirstine et al. 1998). During
harvesting, high quantities of green alcohols and aldehydes are emitted, but isoprene and monoterpenes are
not present (Karl et al. 2001). Green alcohols and aldehydes are considerably less reactive in the atmosphereand do not contribute significantly to O
­ 3 formation.
Monoterpenes were reported to be emitted from somefruit trees including peach (1.2 µg ­g−1 ­h−1) and orange(2.5 µg ­g−1 ­h−1) (Gentner et al. 2014). During the flowering stage, the emission rate increased. The cultivation
area of any fruit tree species in Japan is < 40,000 ha.

5 Discussion
5.1 Interspecific variation in terpenoid emission rate

Figure 2 shows the BERs of isoprene and total monoterpenes from major plant species in Japan. The BERs on aleaf area basis vary widely among plant species. When
comparing data across species, it is observed that the isoprene emission rate is higher than that of total monoterpenes. Two each of deciduous Quercus and bamboo
species are strong isoprene emitters (Fig. 2, upper panel).
Among the monoterpene emitters, four evergreen broadleaf trees, including three Quercus species, have thehighest BERs. The monoterpene storage type conifers L.
kaempferi and P. densiflora have relatively low BERs (0.7–
2.5 nmol ­m−2 ­s−1) that were expressed on the ground areabasis (i.e., tower flux data). These values are similar to themonoterpene fluxes of other coniferous trees grown inEurope and the USA (Tani and Mochizuki 2021).
Emission data on a dry weight basis are summarized forvarious major species and are shown in the lower panelof Fig. 2. Emission data hidden by dark gray bars werecollected using the least reliable SC method and are notdiscussed here. Although most of the data were sourced
from reports distinct from those providing leaf areabased emission rates, a consistent trend was observed.
Specifically, deciduous Quercus species and evergreen
Quercus species exhibited higher emission rates of isoprene and monoterpenes, respectively.
5.2 Terpenoid emission rates within tree species

The leaf area-based BERs of total monoterpenes fromQ. phillyraeoides exhibited up to a 3.1-fold difference
between reports (upper panel of Fig. 2). Similarly, thedifference in the isoprene BER of P. pubescens is 1.8-foldbetween the two reports (Chang et al. 2021b, 2022a). In
contrast, the BERs of Quercus mongolica var. crispulaand Quercus serrata remained consistent at 27.3–27.9
and 27.8–31.7 nmol ­m−2 ­s−1, respectively. These substantial variations in terpenoid emission rates might becaused by factors such as the reliability of measurement
and analytical systems, tree age, leaf morphology, environmental conditions, and genetic diversity within theTani et al. Progress in Earth and Planetary Science(2024) 11:42

Page 14 of 18

Fig. 2 Basal emission rates (BERs) of isoprene and total monoterpenes from major plant species in Japan. Upper and lower panels show BERs
on leaf area basis and dry weight basis, respectively. Dots show the BERs collected in summer season in individual reports. *Expressed on a groundarea basis, **one of three data is expressed on ground area basis. Data hidden by dark bars were removed because they were collected using
a less reliable measurement method (static chamber method). The detailed information including references, measurement techniques, and tree
age for each data is provided in supporting information S1-S3 and Table 3

species. Leaf morphology, particularly leaf thickness,
is linked to cell volume or the number of cells existing
per unit leaf area, which can significantly influence terpenoid production, as it occurs within chloroplasts. For
instance, this phenomenon is evident in isoprene emission from P. pubescens (Chang et al. 2022a). Moreover,various environmental conditions affect terpenoid emissions from plants. For example, monoterpene emissions

from sunlit leaves of Quercus ilex were an order of magnitude higher than those from shaded leaves (Staudt et al.
2003). Furthermore, the light and temperature conditions
experienced by plants during the preceding several to
14 days have been reported to regulate isoprene emission
potentially through enhanced enzyme activity and/or
increased precursor content (Hanson and Sharkey 2001;
Pétron et al. 2001). Genetic diversity may contribute toTani et al. Progress in Earth and Planetary Science(2024) 11:42

intraspecies differences in terpenoid emission rates (Funket al. 2005). The monoterpene emission rate of Q. phillyraeoides was significantly different among 56 individuals,
with six non-emitter variants identified among the treegroup (Chang et al. 2024).
On a leaf dry weight basis, the BER within species varied more than threefold in some plant species, including
Q. serrata, C. japonica, and Q. phillyraeoides (lower panelof Fig. 2). Leaf morphology need not be considered forthe emission rate expressed on a leaf dry weight basis
because leaf weight reflects leaf thickness. Therefore, thevariabilities in terpenoid emission rates within species
may be influenced by the reliability of measurement andanalytical systems, tree age, environmental conditions,and genetic diversity within the species.
5.3 Inventory data required for evaluating the formationof urban O3 and SOA in Japan

Gathering information on the BERs of the top 20 ranked
trees is essential to generate a BVOC inventory in Japanas they cover approximately 70% of the forest area. For
the remaining 30%, limited data are available. The BVOC
inventory will be used to estimate O­ 3 and SOA formations in urban areas to evaluate air quality in current andfuture climates. For this purpose, trees occurring in thealpine and northern areas, including Abies sachalinensis, Betula ermanii, Tilia japonica, and Alnus spp., which
are in the top 20 rankings, are not important. Amongthe other tree species within the top 20, the BER was
not available for Castanopsis cuspidata. The seasonalityof the BER is unavailable for Castanopsis cuspidata var.
sieboldii, C. crenata, Quercus glauca, and C. cuspidata.
Therefore, collecting such data is a priority.
In this review, we describe the high variability in theisoprene and monoterpene emission rates among tree
species. Higher isoprene emission rates were observed in
two deciduous Quercus species and two bamboo species,
and higher monoterpene emission rates were observed inseveral evergreen broadleaf tree species. Among them,
the bamboo species P. pubescens is spreading to suburban areas, including undeveloped woodlands near populated areas known as Satoyama (Takano et al. 2017). The
expansion of invasive bamboo forests can result in an
increase in isoprene emission in suburban areas, whichmight deteriorate air quality around the area.
Large variations in terpenoid emissions have been documented even within the same species. Several factors
contribute to these discrepancies. To obtain representative values of BERs of individual tree species, emission
data must be collected using reliable measurement methods and under representative environmental conditions.
In addition, the average emission rates must be determined using multiple reliable data points.

Page 15 of 18

6 Conclusions
Available data of BERs of isoprene and monoterpenes for
major tree species in Japan were collected from literatureand its datasets were listed in tables. Reliability and validity of the terpenoid measurement methods were evaluated based on their advantages and disadvantages, andthe least reliable data were removed from the datasets indiscussion section. By examining the remaining data, wefound the high variability in the isoprene and monoterpene emission rates among tree species. Higher isopreneemission rates were observed in two deciduous Quercus
species and two bamboo species, and higher monoterpene emission rates were observed in several evergreen
broadleaf tree species. We also found large variationsin terpenoid emissions within the same species. Several
factors may contribute to these discrepancies, including
the reliability of measurement and analytical systems,
tree age, leaf morphology, environmental conditions, andgenetic diversity within the species. To obtain representative values of BERs of individual tree species, emission
data must be collected using reliable measurement methods and under representative environmental conditions.
The BERs need to be determined using multiple reliabledata points.
Abbreviations
BVOCs	Biogenic volatile organic compounds
AVOCs	Anthropogenic volatile organic compounds
O3	Ozone
SOA	Secondary organic aerosols
MEGAN	Model of Emissions of Gases and Aerosols from Nature
PPFD	Photosynthetic photon flux density
BER	Basal emission rate
LAI	Leaf area index
SLW	Specific leaf weight
EAGRID2000-Japan	East Asian Air Pollutant Emissions Grid Database
DMAPP	Dimethylallyl diphosphate
GPP	Geranyl diphosphate
EC	Eddy covariance
REA	Relaxed eddy accumulation
GF	Gradient flux
PTR-MS	Proton transfer reaction mass spectrometryBE	Bag enclosure
LC	Leaf cuvette
FPC	Flow-through plant chamber
SC	Static chamber
LV	Leaf-tip vial

Supplementary Information
The online version contains supplementary material available at https://​doi.​
org/​10.​1186/​s40645-​024-​00645-8.
Additional file 1.
Additional file 2.
Acknowledgements
We thank many our colleagues, particularly K. Endo and K. Inoue, who havegiven helpful comments. We are very grateful to the editor and the two anon‑
ymous reviewers who offered significant improvements to our manuscript.

Tani et al. Progress in Earth and Planetary Science(2024) 11:42

Author contributions
Conceptualization was performed by A.T.; validation by A.T.; data collection by
N.M., T.-W.C., and M.O.; data curation by A.T.; writing—original draft prepara‑tion—by A.T., T.-W.C., N.M., and Y.K.; writing—review and editing—by A.T.;
visualization by N.M., T.-W.C., and A.T.; supervision by A.T.; project administra‑tion by A.T.; funding acquisition by A.T. All authors read and approved the final
manuscript.
Funding
This work was funded by JSPS KAKENHI, grant Number 19H04257 and
19H05666.
Availability of data and materials
The datasets supporting the conclusions of this article are included within thearticle and its additional files.

Declarations
Competing interests
The authors declare that they have no competing interest.
Author details
1
Department of Environmental Sciences, School of Food and Nutritional
Sciences, University of Shizuoka, 52‑1, Yada, Suruga‑ku, Shizuoka 422‑8526,
Japan. 2 Research Institute of Environment, Agriculture and Fisheries, 442
Shakudo, Habikino, Osaka Prefecture 583‑0862, Japan. 3 Tokyo Metropolitan
Research Institute for Environmental Protection, 1‑7‑5, Shinsuna, Kotoku,
Tokyo 136‑0075, Japan.
Received: 4 March 2024 Accepted: 11 July 2024

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