Jurnal Riset Biologi dan Aplikasinya, Volume 5, Issue 1, March 2023
Leaf and Stomata Morphometrics of Gayam Inocarpus fagifer (Fabaceae)
at Different Altitudes
Alwi Smith, Kristin Sangur*, Dessy Fitri Molle, Ludia Haurissa, Grisendy Maulany, Belsefren Renyaan
Biology Education Study Program, Faculty of Teacher Training and Education, Universitas Pattimura,
Jln. Ir. M. Putuhena, Poka Campus, Ambon City, Maluku, Indonesia
*Corresponding Author:
e-mail: sangur_kristin@yahoo.com
Article History ABSTRACT
Received : 5 Desember 2022 Gayam (Inocarpus fagifer) is one of the members of the angiosperm flora in Ambon
City, Indonesia, that grows and develops at various altitudes. This research aimed to
analyze the leaf and stomata morphometrics of these plants in the Aer Louw and
Ema Village areas. Leaf samples were taken from the upper, middle, and lower strata
and considered as replicates. The morphometric characteristics were measured
manually using millimeter block paper and the formula for calculating leaf ratio.
Furthermore, the stomata were stained using the direct incision method and
safranin. The incision results were analyzed using an Olympus CX23 microscope at
400x magnification. The measurement and observation were analyzed descriptively
and correlatively. The results showed that the average leaf width and length, also
the midrib length were greater in Aer Louw Village than in Ema Village; while the
leaf tip and stalk length were greater in Ema Village than in Aer Louw Village. The
characteristics of stomata length and width in Ema Village were greater than in Aer
Louw Village; otherwise, the number, index, and density of stomata in Aer Louw
Village were greater than in Ema Village. Meanwhile, the correlational analysis
showed that the environment influenced the variations of leaves and stomata.
Therefore, the variations of leaves and stomata in the areas could predict plant
adaptations to different environments.
Revised : 12 February 2023
Approved : 22 March 2023
Published : 31 March 2023
Keywords
Altitudinal gradient; ecology;
functional morphology; stomata
density
How to cite: Smith, A., Sangur, K., Molle, D.F., Haurissa, L., Maulany, G., & Renyaan, B. (2023). Leaf and Stomata
Morphometrics of Gayam Inocarpus fagifer (Fabaceae) at Different Altitudes. Jurnal Riset Biologi dan Aplikasinya, 5(1): 16-26.
DOI: 10.26740/jrba. v5n1.p.16-26.
INTRODUCTION
Inocarpus fagifer (Parkinson ex Zollinger) Fosberg
is a woody, leguminous plant with a tree habitus
distributed in tropical and subtropical areas. It has a
shallow taproot, while the lateral roots appear on the
soil surface. The tree bark is rough and brown or gray,
while the leaves are oval, arranged alternately, dark
green, and has a rough surface. The flowers are
arranged in clusters on branches, stems, and twigs with
five petals and showing white or yellowish color
variations. The fruit is oval, irregular, and slightly
flattened, while the young fruit is green and turns
orange-brown when ripe. Moreover, the seeds are
white, fibrous, and thin (Setyowati & Wawo, 2015;
Wawo et al., 2011).
I. fagifer is known in different countries under its
local names, thus aila in Papua New Guinea, chataignier
de Tahiti in French Polynesia, ivi in Fiji, and Tahitian
chestnut or Polynesian chestnut in England (Pauku, 2006).
Pauku et al. (2010) also stated that most farmers use I.
fagifer as an agricultural crop. Meanwhile, I. fagifer in
Ambon City is grown by the community but not
cultivated as a crop, such that the seeds that fall on the
ground will grow naturally. The city is one of the areas
with a large distribution of these plants, namely in Ema
and Air Louw Villages. The distribution in the two
villages represents the highlands and lowlands.
Jurnal Riset Biologi dan Aplikasinya
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https://doi.org/10.26740/jrba.v5n1.p16-26
17| Smith et al., Leaf and Stomata Morphometrics of Gayam
According to Hamidah & Fitriani (2018), some plants
can grow well in the lowlands to the highlands hence,
they have a wide distribution. This explanation shows
that I. fagifer is a plant with a wide distribution in these
two villages. The topography and slope of the place and
environmental factors such as light intensity, wind
speed, temperature, and CO2 pressure vary greatly in
high and low areas (Gao et al., 2019; Kofidis &
Bosabalidis, 2008).
The varying conditions of environmental factors
certainly affect the modification and adaptation of plants
in the two villages. One of the characteristics that is easy
to observe is the morphology of plant organs such as
roots, stems, leaves, flowers, fruits, and seeds. However,
the tree has a large trunk diameter and a height of up to 3
m. This makes it difficult to observe and measure some
morphological characteristics. Meanwhile, flowers, fruits,
and seeds are classified as seasonal organs, making
morphological observation difficult. Leaves are one of the
vegetative organs obtained to observe and measure
morphological characteristics. Leaf morphometric
measurement is conducted to determine leaf area, length,
and ratio, which are also very useful for determining
physiological processes. Observation of stomata
characteristics can easily be performed through the leaf
organ. Liu et al. (2020) also stated that plant adaptation
to changes in environmental factors is carried out by
reducing leaf area and increasing the thickness, mesophyll
tissue thickness, and stomata density.
According to Ruszala et al. (2011) and He & Liang
(2018), stomata are tissues that are very sensitive to the
environment. Therefore, it is very important to observe
and measure the stomata of the I. fagifer leaves to
determine their shape, length, width, aperture, density,
and index. Paembonan et al. (2021) stated that highlands
affect the number of stomata but reduce the size and index
of the Makassar ebony (Diospyros celebica Bakh.). Tumpa
et al. (2022) also noted that geographical location affects
the leaf size of Salix triandra L., a process of
morphological adaptation to environmental changes.
According to Muradoglu & Gundogdu (2011), leaf
surface area relates to the stomata frequency in walnut
plants. Based on these results, the leaf and stomata
morphometric measurement of I. fagifer plants was
conducted based on the difference in altitude in two areas,
namely Ema and Aer Louw Villages. Morphometric
analysis of leaf and stomata at different altitudes can be a
prediction for I. fagifer plants to cope to climate change in
the future. Therefore, this research analyzed leaf and
stomata morphometrics based on different altitudes.
MATERIALS AND METHODS
Sampling Location
This research was conducted in Aer Louw Village
with an altitude 200 m above sea level (asl) and Ema
Village with an altitude of 600 m above sea level (asl)
(Figure 1).
Figure 1. A. Map of Ambon Island; B. Research Locations in Aer Louw Village; C. Research Locations in Ema
Village. Source: https://earth.google.com/web/search/
A
B C
https://earth.google.com/web/search/
Jurnal Riset Biologi dan Aplikasinya, 5(1): 16-26, March 2023 |18
Sampling
Leaf samples for stomata observation and
morphometric measurement were taken separately.
The samples for observing stomata were taken
from one of the largest and tallest I. fagifer trees at
the two sampling locations. Meanwhile,
morphometric measurements were taken from 10
trees in the two locations by considering the upper,
middle, and lower strata. Sampling for stomata
measurement was carried out on the left and right
branches and focused only on green leaves. Leaf
samplings were repeated 5 times with a total was
30 dark green leaves.
Research Procedure
Environmental factors such as temperature,
light, and altitude were measured. The
morphometric measurement of I. fagifer leaves was
conducted in the following stages: (1) leaf samples
were cleaned of dirt and dust using a tissue, (2) the
samples were placed on millimeter block paper and
marked using a pen, (3) the results of the markers
were measured using a ruler as shown in Figure 2.
After obtaining the values for the length, width, tip,
stalk, and leaf midrib length, the next step was to
calculate the formula for the ratio of leaf length and
width (Shi et al., 2020).
The I. fagifer leaf stomata morphometric
measurement was conducted in the following
stages: (1) the leaf samples taken were cleaned of
dirt and dust; (2) the samples were sliced crosswise
at the bottom using a razor blade; (3) the leaf slices
were soaked in commercial bleach (bayclin) for ± 5
min until they turn white; (4) the leaf slices were
washed using distilled water and soaked in 1%
safranin for 1 min; (5) the slices were washed again
using distilled water; (6) the slices were observed
using an Olympus CX23 microscope with 400x
magnification; (7) the observation results were
photographed using an digital camera connected to
a computer; (8) the observed photos were inserted
into the image master to measure length, width and
opening size of stomata, count the number, and
observe the location and type of stomata.
Figure 2. Morphometric characteristics of I. fagifer leaf measurement. (a) leaf length; (b) leaf width; (c) leaf
midrib length; (d) leaf tip length; (e) leaf stalk length
Ratio of leaf length and width =
leaf length
leaf width
Ratio of leaf length and leaf midrib length =
leaf length
leaf midrib length
Ratio of leaf length and leaf stalk length =
leaf length
leaf stalk length
Ratio of leaf length and leaf tip length =
leaf length
leaf tip length
19| Smith et al., Leaf and Stomata Morphometrics of Gayam
Figure 3. Morphometric characteristics of I. fagifer leaf stomata measurement. (a) stomata length; (b) stomata
width; (c) stomata opening size
Data Analysis
Data from I. fagifer leaf and stomata
morphometric calculation were collected and
analyzed descriptively to determine the average
value and standard deviation. Furthermore, the data
were analyzed to determine correlation value of
environmental factors and effective contribution
using multiple linear regression inferential statistics
(Wang et al., 2019; Sun et al., 2021). An analysis of
correlation value and effective contribution is used
based on the following formula:
Formula of product moment correlation
𝑟𝑥𝑦 =
𝑛 ∑ 𝑋𝑌−(∑ 𝑋) (∑ 𝑌)
√{𝑛 ∑ 𝑋2− (∑ 𝑋)2}
(Chawla et al., 2016; Kumari & Yadav, 2018).
Then analyze the effective contribution (EC):
𝐸𝐶%𝑋𝑛 = 𝐵𝑋𝑛 𝑥 𝑟𝑥𝑦 𝑥 100%
(Turkheimer & Waldron 2000).
Note:
EC: effective contribution; BXn: B coefficient of the
predictor; Xn: predictors such as temperature, light,
and altitude, rxy: correlation coefficient.
The calculation for the stomata index and
density was conducted based on the Fetter et al.
(2019) formula as follows:
Stomata Density =
Number of Stomata
Field of View Unit
Index Stomata =
Number of stomata
Number of stomata + epidermal cells
The data was analyzed using Excel and SPSS for
Windows 18.
RESULTS AND DISCUSSION
Inocarpus fagifer Leaf and Stomata Morphology
Morphologically, the color of I. fagifer leaves in
Ema and Aer Louw Villages was the same, dark
green on the upper surface and light green on the
lower surface. The upper surface of the leaves is
smooth and greasy, while the lower surface was
rough (Figure 4). Meanwhile, the stomata
morphology between the two areas has the same
shape but differed in size and number of stomata in
one field of view (Figure 4).
Stomata are a type of differentiation from leaf
epidermal tissue (Peterson et al., 2013; Torii, 2021;
Zuch et al., 2022). I. fagifer stomata are found on the
lower surface of the leaves. The stomata in these two
areas have an actinostic type, with guard cells
surrounded by neighboring cells in a radius. The
number of neighboring cells is 4 or more, while the
stomata’s guard cells are kidney-shaped with thin
side walls and thicker top and bottom walls
(Prabhakar, 2004; Ahmad et al., 2009; Song et al.,
2020).
Jurnal Riset Biologi dan Aplikasinya, 5(1): 16-26, March 2023 |20
Figure 4. I. fagifer leaf morphology. (Top row) Samples from Ema Village (Altitude of 600 m asl). (Bottom
row) Samples from Aer Louw Village (Altitude of 200 m asl).
Leaf and Stomata Morphometrics of Inocarpus
fagifer
The morphometric measurement in the two
areas with different altitudes varies greatly, as
summarized in Table 1. The same result was
reported by Paridari et al. (2013) wherein Carpinus
betulus L. growing at high altitudes had a small leaf
lamina compared to those growing at low altitudes.
According to Liu et al. (2020), adapting plants in
the highlands reduces leaf area. This shows that I.
fagifer plants that grow at different altitudes have
adapted to have an average leaf width and length at
high altitudes (Ema Village) of 119.35 mm and
259.77 mm, while those growing at low altitudes
(Aer Louw Village) have an average leaf width and
length of 135.43 mm and 312.52 mm. The long and
wide leaves of the I. fagifer plants in Aer Louw
Village also have a midrib average length of 268.47
mm, while in Ema Village the midrib average
length is 245.8 mm. This result agrees with that of
Madeline et al. (2014), where broad leaves have
high vein density accompanied by stomata density.
Meanwhile, the broad leaves of I. fagifer in Aer
Louw Village had shorter tips and petioles on
average 3.69 mm and 5.45 mm, while the average
tips and petioles in Ema Village were 6.07 and 8.7.
According to Serdar & Kurt (2011), leaf parameters
can be used as a variable to detect the level of
phenotypic variability among plant species in a
population. The stomata morphometric
measurement of I. fagifer leaves in the two areas with
different altitudes varied greatly, as recapitulated in
Table 2. The high altitude in Ema Village (600 m asl)
resulted a lower stomata density than Aer Louw in
the lower area (200 m asl). Fustier et al. (2019)
reported that stomata density decreased with
increasing altitude. According to Li et al. (2021),
plants with large stomata have low densities, but
large size affects plant adaptation. Furthermore, Idris
et al. (2019) stated that high intensity affects stomata
density to support high assimilation processes in
plants.
21| Smith et al., Leaf and Stomata Morphometrics of Gayam
Table 1. Morphometric characteristics of I. fagifer leaves at two places with different altitudes
Morphometric
Characteristics of I. fagifer
Leaves
Location
Upper strata
(mm)
Middle strata
(mm)
Lower strata
(mm)
Leaf length
Ema 265.7±3.37 258.2±4.03 255.4±4.17
Aer Louw 302.26±2.12 293.22±1.03 342.08±18.4
Leaf width
Ema 116±2.93 111.8±2.63 111.7±2.83
Aer Louw 140.14±2.12 135.88±1.03 130.26±0.82
Leaf midrib length
Ema 251 ±3.22 244.7±3.83 241.7±3.92
Aer Louw 278.26±4.23 269.22±2.89 257.94±2.37
Leaf tip length
Ema 9.7±0.17 4.1±0.12 4.4±0.14
Aer Louw 3.66±0.66 3.76±0.04 3.66±0.05
Leaf stalk length
Ema 8.8±0.26 8.7±0.26 8.6±0.3
Aer Louw 5.40±0.1 5.56±0.07 5.40±0.1
Ratio of leaf length and width
Ema 23.2±0.37 23.09±21.56 22.8±0.29
Aer Louw 21.64±0.19 21.56±0.14 25.56±1.16
Ratio of leaf length and leaf
midrib length
Ema 10±0 10±0 10±0
Aer Louw 10.99±0.12 10.89±0.05 13.09±0.64
Ratio of leaf and leaf stalk
length
Ema 303±7.93 294.5±6.83 314.4±6.28
Aer Louw 487.9±27.9 500.12±18 535.68±54.2
Ratio of leaf length and leaf tip
length
Ema 535.1±32.3 59.04±12.3 531.7±15.6
Aer Louw 856.5±24.3 785.01±9.19 966.59±68
Table 2. Stomata morphometric characteristics of I. fagifer leaves at two areas with different altitudes
Stomata
Morphometric
Characteristics
Loc.
Upper Strata (μm) Middle Strata (μm) Lower Strata (μm) Desc
riptio
n
Right
Branch
Left
Branch
Right
Branch
Left
Branch
Right
Branch
Left Branch
Stomata Length
(Mean±SD)
Ema 47.54±
0.40
47.50±
0.33
44.07±
0.26
44.21±
0.45
41.88±
0.39
42.39±
0.46
Very
long
Aer
Louw
16.68±
0.14
16.63±
0.12
14.89±
0.22
14.96±
0.21
12.75±
0.995
12.79±
0.75
Less
long
Stomata Width
(Mean±SD)
Ema 46.86±
0.56
47.09±
0.39
44.4±
0.26
44.02±
0.37
42±
0.46
41.99±
0.63
Very
wide
Aer
Louw
16.93±
0.30
17.09±
0.29
14.96±
1.06
15.26±
0.54
14±
1.07
14.3±
1.64
Less
wide
Stomata
Opening Size
(Mean±SD)
Ema 16.03±
0.27
16±
0.32
14.11±
0.34
14.06±
0.25
12±
0.24
12.02±
0.24
Wide
Aer
Louw
6.058±
0.11
6.2±
0.13
5.01±
0.1
5.16±
0.24
4.8±
0.39
4.92±
0.46
Less
wide
Number of
Stomata
(Mean±SD)
Ema 13±
1.52
14.4±
2.30
10±
0.71
9.8±
0.84
7.2±
0.84
6.4±
1.14
Few
Aer
Low
43±
7.01
40.2±
9.15
24.4±
4.62
27.4±
8.05
24±
3.27
21.4±
1.52
Many
Stomata Index
(Mean±SD)
Ema 0.18±
0.01
0.18±
0.01
0.15±
0.01
0.15±
0.01
0.1±
0.02
0.1±
0.01
Low
Aer
Louw
0.58±
0.02
0.57±
0.03
0.43±
0.02
0.44±
0.01
0.37±
0.01
0.36±
0.01
High
Stomata
Density
(Mean±SD)
Ema 152.3±
17.24
163.6±
26.16
113.6±
8.04
111.4±
9.51
81.81±
9.51
72.72±
12.96
Low
Aer
Louw
670.8±
108.9
624.2±
142.1
379.9±
70.23
425.5±
124.9
370.6±
48.9
333.3±
21.61
High
Description: Loc: Location
Jurnal Riset Biologi dan Aplikasinya, 5(1): 16-26, March 2023 |22
The difference in altitude is an environmental
factor affecting the plant microclimate. According to
Lamprecht et al. (2018), ecosystems at high altitudes
have low temperatures. Meanwhile, Idris et al. (2019)
reported that the stomata density increased when
exposed to high sunlight. Environmental
characteristics at different altitudes also affect the
stomata morphometric features. Tiwari et al. (2013),
also stated that altitude was positively correlated
with stomata density, index, and guard cell length.
According to Akbarinia et al. (2011), variations in
shape, size, index, area, and stomata can vary within
one species. The stomata length characteristic of I.
fagifer leaves is directly proportional to its width.
Muradoglu & Gundogdu (2011) also stated a positive
relationship between stomata length and width.
According to Li et al. (2011), the stomata index of
Quercus aquifolioides Rehder & E.H. Wilson decreased
at high altitudes and increased at low altitudes.
Meanwhile, the morphological characteristics of
stomata related to its density are inversely
proportional to the length and width of it, as well as
to the size of the stomatal opening, which is inversely
proportional to stomata density (Hong et al., 2018;
Haworth et al., 2023).
Some of these findings have supported this
research that the length of the I. fagifer stomata
leaves are also directly proportional to the width of
the stomata and the size of the opening of the
stomata is directly proportional to the number,
index, and density of stomata.
The variation of stomata in the two areas with
different altitudes shows that altitude plays a role in
morphometric characteristics. According to Alonso-
Amelot (2008), highland plants have high
adaptability to extreme environments. It was stated
by Ahmad et al. (2020) that the ability of plants to
adapt in the highlands is by adjusting their
morphological and physiological characteristics.
Halbritter et al. (2018) and Montesinos‐Navarro et
al. (2011) also confirmed that the elevation gradient
greatly affects abiotic factors, such as humidity,
temperature, and light intensity in an area.
Variations in leaf and stomata morphometrics of I.
fagifer as affected by environmental parameters
Environmental characteristics in the two areas with
different altitudes are shown in Table 3. The condition
of the two areas showed that light intensity
influences temperature, while altitude is related to
light intensity as indicated in Table 3. The condition
of the two areas showed that light intensity
influences temperature, while altitude is related to
light intensity as indicated in Table 3. Altitude is an
environmental factor that greatly determines the
relationship between leaf and stomata
morphometrics in I. fagifer plants. The relationship
of environmental factors to the leaf morphometric
characteristics of I. fagifer plants is shown in Table 4.
Table 3. Environmental characteristics
Environmental Characteristics Ema Village Air Louw Village
Light intensity 17,000 Lux 20,000 Lux
Temperature (ᵒC) 25ºC 28ºC
Altitude 600 m asl 200 m asl
Table 4. Correlation of environmental factors with leaf morphometric characteristics
Leaf
Characteristics R R
2
Sum of Square Mean Square
F F sig (p)
Reg. Res. Reg. Res.
Leaf length 0.31 0.099 417.4371 3798.868 417.43713 65.5 6.3733
08
0.014(*)
Leaf width 0.46 0.2136 74.32614 273.564 74.32614 4.717 15.758
35
0.00(*)
Leaf midrib
length
0.32 0.1025 77.11201 675.319 77.112007 11.64 6.6227
93
0.012(*)
Leaf tip length 0.32 0.1003 0.074907 0.671787 0.074907 0.012 6.4672
12
0.014(*)
Leaf stalk length 0.65 0.418 1.581127 2.201347 1.581127 0.038 41.658
75
0.00(*)
Ratio of leaf
length and width
0.01 0.0001 0.001815 16.26631 0.001815 0.28 0.0064
72
0.936
23| Smith et al., Leaf and Stomata Morphometrics of Gayam
Leaf
Characteristics R R
2
Sum of Square Mean Square
F F sig (p)
Reg. Res. Reg. Res.
Ratio of leaf
length and leaf
midrib length
0.3 0.09 0.411682 4.163537 0.411682 0.072 5.7349
17
0.019(*)
Ratio of leaf
length and leaf
stalk length
0.42 0.1727 7390.602 35402.1997
6
7390.6021 610.4 12.108
14
0.001(*)
Ratio of leaf
length and leaf
tip length
0.47 0.2163 16625.69 60222.5031
4
16625.692 1038 16.012
12
0.00(*)
Description: Reg: Regression; Res: Residual; (*): significant
Table 5. Effective contribution of environmental factors to leaf morphometric characteristics
Leaf characteristics
Effective Contribution of Environmental
Factors Total (%)
Temperature Light Altitude
Leaf length 0.00 0.00 9.901 9.901
Leaf width 0.00 0.00 21.36 21.36
Leaf midrib length 0.00 0.00 10.25 10.25
Leaf tip length 0.00 0.00
10.03 10.03
Leaf stalk length 0.00 0.00
41.8 41.8
Ratio of leaf length and width 0.00 0.00
- -
Ratio of leaf length and leaf midrib length 0.00 0.00 8.998 8.998
Ratio of leaf length and leaf stalk length 0.00 0.00 17.27 17.27
Ratio of leaf length and leaf tip length 0.00 0.00 21.63 21.63
Table 6. Correlation of environmental factors with stomata morphometric characteristics
Stomata
Characteristics
R R2
Sum of square Mean square
F F sig (p)
Reg. Res. Reg. Res.
Stomata length 0.992 0.983 2667.697 44.952 2667.697 4.495 593.452 0.00(*)
Stomata width 0.992 0.985 2493.795 39.027 2493.795 3.903 638.997 0.00(*)
Stomata opening
size
0.964 0.93 229.338 17.248 229.338 1.725 132.961 0.00(*)
Number of stomata 0.843 0.711 1180.083 480.833 1180.083 48.08 24.542 0.00(*)
Stomata index 0.915 0.838 0.288 0.056 0.288 0.006 51.575 0.00(*)
Stomata density 0.879 0.772 370572.395 109402.096 370572.4 10940 33.873 0.00(*)
Description: Reg: Regression; Res: Residual; (*): significant
Table 7. Effective contribution of environmental factors to stomata morphometric characteristics
Stomata characteristics
Effective Contribution of Environmental Factors Total
(%) Temperature Light Altitude
Stomata length 0.00 0.00 98.3 98.3
Stomata width 0.00 0.00 98.5 98.5
Stomata opening size 0.00 0.00 93 93
Number of stomata 0.00 0.00 71.1 71.1
Stomata index 0.00 0.00 83.8 83.8
Stomata density 0.00 0.00 77.2 77.2
Jurnal Riset Biologi dan Aplikasinya, 5(1): 16-26, March 2023 |24
The relationship of environmental factors to the
stomata morphometric characteristics of I. fagifer
leaves is shown in Table 6. Environmental factors of
light, temperature, and altitude have a significant
relationship with all morphometric characteristics of
I. fagifer leaves (p=<0.05). Previous research
confirmed that environmental factors greatly affect
stomata opening size (Casson & Gray, 2008). Qi &
Torii (2018), reported that environmental factors
stimulate stomata density. Harrison et al. (2020) also
stated that environmental factors correlated with
stomata size and density.
The effective contribution of environmental
factors was calculated to determine which stomatal
morphometric characteristics were more dominant.
The altitude effectively contributed to these stomatal
morphometric characteristics, as indicated in Table
7. Aslantaş & Karakurt (2009) stated that high areas
have high rainfall while temperature, O2 and CO2
levels decreased. This shows that the environmental
factors of temperature, light, O2, CO2, and humidity
depend on altitude.
The low and high altitudes are related to
temperature, light, and humidity. These
environmental factors affect stomata length, width,
opening size, number, index, and density
simultaneously. Specifically, stomata opening is
influenced by light (Elhaddad et al., 2014) and high
temperature (Lawson & Blatt, 2014). Driesen et al.
(2020) stated that stomata opening is influenced
simultaneously by light, CO2, temperature, and
humidity. Altitude greatly influences plant
physiology, such as stomata density (Qiang et al.,
2003). Richardson et al. (2017) confirmed that
stomata are adaptive tissues that modify their
stomatal density, size, and form in response to
environmental changes.
CONCLUSION
The results showed that different
environmental conditions can provide variations in
the morphology of the leaves and stomata of I. fagifer
plants. Altitude is related to other environmental
factors, such as temperature and light intensity,
which can directly influence variations of leaves and
stomata. This research can predict I. fagifer plants’
survival and adaptation to environment changes.
ACKNOWLEDGEMENT
The authors are grateful to the leadership of the
Faculty of Teacher Training and Education at
Pattimura University, which has provided funds for
this research. The funding is stated in the Certificate
Number 1087/UN13/SK/2021.
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