International Journal of Environment, Agriculture and Biotechnology

Vol-7, Issue-6; Nov-Dec, 2022

Journal Home Page Available: https://ijeab.com/

Journal DOI: 10.22161/ijeab

Peer Reviewed

Evaluation of the cross-pollination in maize (Zea mays L.)

synthetic varieties grown in the High Guinean savannah
zone conditions
Maygon Katoukam1,*, Maina Antoine Nassourou2, Souina Dolinassou2, Jean-Baptiste
Tchiagam Noubissié1
1
Department of Biological Sciences, Faculty of Sciences, University of Ngaoundéré, P.O. Box 454 Ngaoundéré, Cameroon
2Department of Biological Sciences, Faculty of Sciences, University of Maroua, P.O. Box 814 Maroua, Cameroon
*Corresponding Author

Received: 20 Nov 2022; Received in revised form: 13 Dec 2022; Accepted: 22 Dec 2022; Available online: 29 Dec 2022
©2022 The Author(s). Published by Infogain Publication. This is an open access article under the CC BY license
(https://creativecommons.org/licenses/by/4.0/).

Abstract— In Cameroon, maize is the most widely cultivated cereal and is consumed by more than one third
of the population. This study aimed to evaluate in the tropical conditions the cross-pollination rate in four
recipient synthetic maize varieties by xenia phenomenon depending on distance to the pollen source and
wind direction. The experimental design was triplicated split plot with each replication arranged as a 576
m2 Latin square area. The combined analysis of variance showed a highly significant effect (p<0.001) of the
gap from pollen source and wind direction on the cross-pollination rate of the recipient varieties. CMS 8704
yellow-grain variety which is the pollen donor and the white grain receiver cultivars CMS 2019, CMS 8501,
CMS 9015 and Shaba had one to seven days’ difference between the female flowering of the recipient variety
and the start of male flowering of the donor. These synthetic varieties differed significantly for the number
of leaves per plant, the 100-seeds weight, the plant height, and total kernels weight per plant, with cultivar
Shaba showed the highest values. The highest cross-pollination rates were found in the first maize rows
facing the donor field and the genetic pollution decreased with increasing distance from the donor source.
At the same distance from source, the pollution level higher the North. The implementation of appropriate
separation distance (>10 m) is recommended for reducing genetic pollution and ensuring coexistence of
different genotypes in maize production field.
Keywords— Cameroon, Cross-pollination, Genetic pollution, High Guinean savannah, Maize.

I. INTRODUCTION plays a prominent role in the advent of maize across the

Maize (Zea mays L., 2n=20.) is the most cultivated global agri-food system (Brouwer et al., 2020). Genetically
plant in the world and the first cereal that enters in the diet modified (GM) maize is the second most important GM
before wheat (Garcia-Lara and Serna-Saldivar, 2019). It is crop following soybean (Willet et al., 2019). The world’s
a versatile multi-purpose crop, primarily used as a feed total maize production was estimated at 1.05 million
globally, but also is important as a food crop, especially in thousand tons in 2020 (Erenstein et al., 2022). Maize
sub-Saharan Africa and Latin America, besides other non- production of Cameroon was estimated at 2.200 thousand
food uses (Grote et al., 2021). The maize-distilling process tons in 2020 (FAO, 2021). The current production level of
has long been used by industry for the production of maize in the country is declining and to meet consumption
beverage alcohol. Maize thereby plays a diverse and requirements, huge quantities of the commodity are
dynamic role in global agri-food systems and food/nutrition imported (Mvodo Meyo and Mbey Egoh, 2020).
security (Poole et al., 2021). Improved maize germplasm

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Maize is predominantly cross-pollinated with vigor as well as seed yield and its components (Denney,
anemophily as the general rule. Maize is a wind-pollinating 1992). Understanding pollen mediated gene flow is also
crop with about 95% cross pollination (Devos et al., 2007). important to achieve the coexistence measures for farming
Pollination with insects also takes place to certain extent with and without genetically modified and conventional
(Klein et al., 2007). The adaptation for cross-pollination are maize (Messeguer et al., 2006; Njountie Tchiengue, 2010).
monoecious inflorescences, unisexual flowers, differences Cross-pollination is affected by many factors inducing
in time of maturity of the male and female inflorescence, distance between donor and recipient fields, wind direction,
silk receptive on entire length and abundant pollen wind speed, flowering synchronization between donor and
production (Brittan, 2006). The intensification of maize recipient plants, field topography, size and orientation of
production in order to reduce the food deficit in the face of donor and recipient fields, pollen velocity, weather
galloping demographics requires an acceleration of the condition like any temperature and air humidity (Devos et
creation of new, better performing and better adapted al., 2005; Vogler et al., 2009). Most of pollen settles within
varieties. Hybrid maize requires new materials for every 06 to 15 m of the donor plant (Brittan, 2006). The main
crop to maintain its potential and proved a particularly purpose of the present study was to evaluate on some
viable and attractive business model for the seed industry synthetic maize varieties grown in Dang (Adamawa-
(Morris et al., 2003). In sub-Saharan Africa, the Cameroon) the pollen mediated gene flow depending on
development of improved open pollinated varieties (OPVs) source-recipient distance and wind direction.
and synthetic varieties were recommended to smallholder
farmers for their performance, and their seed-recycling
II. MATERIAL AND METHODS
potential (Morris et al., 2003). Composite or synthetic seeds
are the most appropriate for developing countries because 2.1 Study area
they give farmers the possibility to renew the seed from The study was carried out from 2020 to 2021 at the
their harvests in addition to their productivity. However, the University of Ngaoundéré experimental farm, at Dang
stability of composite varieties depends in part on the level (Ngaoundéré 3rd subdivision, Adamawa region, Cameroon),
of genetic pollution which is the accidental transfer of genes which is intersected by 7° 26' 16 4'' North latitude and 13°
between genotypes through inter-pollination (Tsai and Tsai, 33' 34'' East longitude and has 1115 m above the mean sea
1990). level. This region belongs to the Guinea High Savannah
Cross-pollination studies between adjacent maize agroecological zone (Djoufack et al., 2012). The climate is
fields have been conducted all over the world using mainly of the Sudano-Guinean type characterized with a humid
a color marker system (Messeguer et al., 2006; Njountie trend, an average annual rainfall of 1480 mm distributed
Tchiengue, 2010). The main focus of these studies was to over the rainy season (March-October), and a dry season
gather information about adequate separation distances to (November-March). The average annual temperature is
ensure coexistence and about the dependence of cross- 22.59°C, while the relative humidity is about 66.47%. The
pollination on the distance within maize fields. Several soil in the area is mostly ferruginous type developed on old
studies have been performed to evaluate the impact of basalt and has a brown reddish clay texture. There is an
pollen drift from fields containing GM corn to neighboring immense dependence of agriculture productivity on soil
non-GM cornfields (Byrne and Fromherz, 2003; Devos et physicochemical properties (Nanganoa et al., 2020).
al., 2005; Weber et al., 2007; Njontie Tchiengue, 2010; 2.2 Plant material
Viorica et al., 2017). The accidental gene flow is more
The plant material used consisted of five composite
pronounced under the conditions of peasant agriculture,
maize cultivars adapted to the Guinea High Savannah
where crop plots are close together and sometimes several
agroecological zone, comprising a yellow grain (CMS
varieties are grown in the same plot. The cultivation of
8704) using as pollen donor and four recipient white grain
maize in Cameroon is predominantly dominated by
(CMS 9015; CMS 8501; CMS 2019; CMS 8806 and Shaba
smallholder farmers who use traditional methods and face
(Table 1). The seeds were obtained from the Institute of
drudgery. The effect of genetic pollution on maize can have
Agricultural Research for Development (IRAD, Garoua
many consequences in particular considerable variation in
station, Cameroon).

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Table 1. Characteristics of the tested synthetic maize varieties.

Varieties Cycle (days) Tasseling Days to female Seed Seed
(days) flowering color texture
CMS 2019 (receiver) 110 -115 58 - 60 61 – 64 White Horned
CMS 8501 (receiver) 105 -110 55 - 58 59 – 62 White Toothed
CMS 8704 (donor) 105 -110 57 - 59 58 – 61 Yellow Horned
CMS 9015 (receiver) 90 – 95 55- 58 58 – 62 White Toothed
SHABA (receiver) 110 – 130 61 - 65 64 – 69 White Toothed

2.3 Field trials index or 100-seeds weight (SI) and the difference between
During the growing season 2020, the sowing was the female flowering on recipient’s varieties and the start of
done simultaneously, for the donor and for the pollen male flowering of the donor. The time difference between
receiver’s varieties. The experiment was laid out in a male and female flowering of a single plant is called
triplicated split plot design consisting of eight source- anthesis-silking interval (ASI). In the case of cross-
recipient distances (main treatment), four sub-treatments pollination, the difference in days between the female
(wind directions), with each replication arranged as a 576 flowering of the recipient variety and the start of male
m2 Latin square area (24.0 m x 24.0 m) (Fig. 1). In the flowering of the donor is ASI2 (Devos et al., 2005). The
experimental field the three blocks were spaced 120.0 m number of leaves per plant was obtained by manual
each other to avoid cross-fertilization. In the center of each counting on the sample of 20 plants per variety randomly
square, CMS 8704 yellow-grain variety which is the pollen selected during the flowering. The height of the plant was
donor was sown inside an area of 16 m 2 (4 m length x 4 m measured using a graduated decameter. The total kernel
broad). Recipients white varieties were sown each on an weight per plant and the 100-seed mass were determined
experimental unit consisting of one row of 4.0 m length, using an electronic balance of 0.001g sensitivity (Sartorius
respectively at 1.5 m, 2.5 m, 4.0 m, 5.0 m, 6.5 m, 7.5 m, 9.0 Prodilab).
m and 10.0 m from the pollen-donor source. Three seeds
were sown per hill and one seedling was retained after 2.2 Evaluation of genetic pollution rate
thinning. Recipient varieties were sown in four different
The donor maize was a yellow grain (dominant
wind orientations (West, East, South and North). Maize
trait) cultivar CMS 8704 and the recipient maize was white
plants were spaced 25.0 cm for receiver’s plots and 40.0 cm
grain (recessive trait) cultivars CMS 2019, CMS 8501,
for donor plots. All recommended agricultural practices
CMS 9015 and Shaba. This would enable us to easily
were adopted throughout the field trials, except the
distinguish intra-cultivar pollinated and inter-cultivar
application of pesticides. NPK (20% N, 10% P2O5, 10%
pollinated grains through xenia phenomenon as
KO2) and urea (46% N) fertilizers were applied to the soil
recommended by Denney (1992). The xenia usually refers
at 20 and 45 days after sowing respectively. Regular manual
to a situation in which the genotype of the pollen donor
weeding was carried out during the vegetative phase and at
influences the maternal tissue of the fruit so as to produce a
flowering. At maturity, a total of 10 plants were randomly
phenotypically demonstrable effect upon the seed grains of
selected in each row of the recipient field for the evaluation
the recipient. When the ovules of the recipient varieties
on cross-pollination rate. On each selected plant, their main
were fertilized with the pollen of the yellow grain variety,
ear was collected, and the kernel number determined by
the grains obtained appeared yellow, thereby displaying the
counting separately white and yellow grains.
xenia effect. Cross-pollination was investigated by the
2.1 Characterization of maize genotypes presence of yellow-grains on white-grain varieties at
The characterization of the five synthetic varieties distance up to 17.5 m from the yellow-grain pollen (Vogler
used in the study was done by randomly selecting 20 plants et al., 2009). By counting the grains showing xenia among
per genotype in each of the replications (five plants for each the total grains per ear of a recipient, we could easily
direction). The experimental design is a triplicated non estimate the cross-pollination rate (P) from the following
randomized complete block design. Four characters formula:
selected among the maize descriptors were retained: the Number of yellow grains
height of the plant (HP), the number of leaves per plant P= × 100
Total number of grains
(NLP), the total kernel weight per plant (KWP), the seeds
With P: the cross-pollination rate in percentage

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2.3 Statistical analysis Student’s t-test at 5% level of probability. Pearson linear

Data obtained were subjected to the analysis of correlation coefficient was used to assess the relationships
variance (ANOVA) using Statgraphics Plus Version 5.0 between cross-pollination rate and distance from the source
software. Differences in means performance were tested of pollen.
using the Least Significant Difference (LSD) or by the

24m

North
East

West
24m

South

Fig. 1: Experimental layout representing pollen receiver varieties around the donor variety according to the four cardinal
points (wind direction).

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III. RESULTS AND DISCUSSION weight per plant of 144 g to 357 g on improved varieties of
3.1 Variability of tested maize varieties maize and noted that the weight of grains could be predicted
on the length of ear based on the linear equation y = 22.5x -
The analysis of variance showed significant
156.9. The 100-seeds weight ranged from 29.11 g for
variability (p<0.05) among the five maize varieties for plant
yellow grain donor variety CMS 8704 to 49.64 g for Shaba
height, number of leaves per plant, seed index and the total
with an average of 35.72 g (Table 2). The seed index values
weight of kernels per plant (Table 2). The average plant
recorded in this study were greater than values recorded by
height varied from 183.0 cm (CMS 8501) to 272.21 cm
Inamullah et al. (2011) on maize hybrids. The variability
(Shaba) with a mean of 207.23 cm. The height the yellow
noted for these traits could be due to the genetic diversity
donor was slightly greater than those of the white receptor
and environmental conditions under which the trials were
plants except Shaba. Similar results were obtained by
conducted. There is genetic variability within cultivars for
Vogler et al. (2009) showing that maize height varied
most of the agro-morphological traits. The creation and
between 219 cm and 250 cm for improved varieties. In
maintenance of growth and developmental homogeneity
contrast, Viorico et al. (2017) in Romania observed that
within maize population is essential.
some improved maize varieties had tallest height.
The synchronization between of the start of male
For the number of leaves, the highest value was
flowering of the donor variety CMS8704 and female
noted for popular Shaba cultivar (16.20 leaves) while the
flowering of the four white grain recipients ranged from one
lowest was 14.74 leaves (CMS 8501). The stem of maize is
to seven days (Table 2). As the synchronization of yellow
commonly composed of 20 internodes and the leaves arise
male flowering and white female flowering was much
from these nodes. The number of leaves per plant noted in
closer than the flowering of white male and female plants
this study at flowering was in agreement of reports of
except for Shaba, cross-pollination could be expected to be
Sangoi and Salvador (1997). The number of leaves per plant
unusually high (Aylor et al., 2003) However, rates of cross-
seemed to increase with the size of the plant. Plant height is
pollination were in the expected range (Ma et al., 2004;
a good indicator to evaluate plant growth and grain yield.
Messeguer et al., 2006; Bannert and Stamp, 2007; Della
The dynamic of plant height during the growing cycle could
Porta et al., 2008). The ASI2 depends on the genotype and
be used to access critical genetic traits, fundamental plant
environmental factors like water deficit, nutrient light and
physiology and environmental effect. The vertical
temperature (Devos et al., 2005). The difference in sowing
distribution of leaf is important for the analysis of
dates may influence the flowering times, and limiting cross-
photosynthesis, stress resistance and pollen propagation.
pollination. Synchronization between donor and receiver of
Total kernel weight per plant varied from 154.14 g pollen is very important for inter-crossing between
(CMS 9015) to 204.18 g for Shaba with an average of varieties.
169.95 g. Drienovsky et al. (2019) recorded a total kernel
Table 2. Genetic variability of some characteristics of tested maize varieties
Variety Parameters
PH (cm) NLP KWP (g) SI (g) ASI2 (days)
CMS 8704 192.88±6.42b 14.93±0.22b 174.33±7.11b 29.11±3.36c -
c b c b
CMS 8501 183.0±4.02 14.74±0.23 156.89±2.30 35.99±3.35 2.0
c ab c bc
CMS 9015 184.33±6.42 15.12±0.17 154.14±4.83 31.05±3.36 1.0
CMS 2019 189.47±5.35bc 14.76±0.48b 166.27±9.11bc 32.81±2.84bc 4.0
a a a a
Shaba 272.21±14.43 16.20±0.29 204.18±6.06 49.64±4.67 7.0
Means 204.37±7.32 15.15±0.27 170.76±5.88 35.72±3.51 3.5
LSD (5%) 8.20 1.01 16.52 5.22

Values followed by the same letter on the line are not significantly different (p<0.05). PH: plant height; NLP: number of leaves
per plant; KWP: total kernels weight per plant; SI (g): Seed index or 100 seeds weight; ASI2: difference in days between the
female flowering of the recipient variety and the start of male flowering of the donor.

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3.2 Effect of the distance from pollen source distance is affected by the height of pollen release, and the
and wind direction on the cross-pollination topography of the surface. From these results, it appeared
rate of recipient maize varieties that at an isolation distance of 10 m, genetic pollution rate
The analysis of variance showed that the cross- was less than 7%. These results demonstrated that spatial
pollination rate of the recipient maize varieties varied isolation is an effective method to reduce outcrossing rates
significantly (p<0.001) with the distance from the pollen in maize.
source and the wind direction (Table 3). The interaction Concerning the wind direction (Table 3, Fig. 3),
between wind direction and distance from pollen source and xenia percentage was highest in north (19.27%) and west
the blocks effects were not significant. These results tell us (17.01%) and decreased in south (13.78%) and east
that allogamy rates depend on the distance from the donor (14.26%). Ma et al. (2004), pointed out that the cross-
pollen source and the wind direction. Raynor et al. (1972) pollination rate was significantly higher downwind than
and Ma et al. (2004) noted that environmental factors and upwind from the pollen source. Weber et al. (2007) noted
distance influenced cross-pollination in maize. The highest that the influence of wind can change between locations and
rates of cross-pollination were found closest to the pollen years, so reliable prediction is not possible. However, wind
source and at further distances from the pollen source the speed and direction cannot be reliably incorporated into
decrease in cross-pollination was much stronger (Table 4, strategies to avoid cross-pollination. Measurements of
Fig. 2). The average pollution rate was 38.6% at 1.5 m from horizontal wind speed during flowering in relation to the
the source and decreased to 5.3% at 10.0 m. Cross- sedimentation rate of maize pollen showed a potential
pollination studies between adjacent maize fields were distance for horizontal pollen dispersal (Bannert and Stamp,
conducted worldwide using mainly a colored marker 2007). The few cross-pollinations observed over longer
system. Many recent studies also noted that xenia distances could be due to gusty or vertical wind movements
percentage was highest at the border rows facing the donor (thermal or turbulence effects). According to Hofmann et al.
and decreased rapidly with increasing distance from the (2014), most corn pollen falls within the first five meters
donor field (Aylor et al., 2003; Ma et al., 2004; Viorica et past the edge of the field, but the possibility of finding a
al., 2017). According to Devos et al. (2005), most pollen small amount of pollen at greater distances from the corn
coming from donor field was retained at the border rows of plot depends on wind speed.
recipient field that constitute a protection band, thus the Cross-pollination depend on other factors. A
proportion of donor pollen within the recipient field will difference in flowering between the donor and recipient can
decrease. The majority of the pollen deposition took place reduce the level of cross-pollination (Devos et al., 2005).
within the first two meters of the pollen source but the Timing between anthesis of the pollen donor and silking of
possibility to find a small amount of pollen at larger the recipient is one of the main factors affecting the pollen-
distances from the source exist. These observations mediated gene. Della Porta et al. (2008) observed that cross-
recorded in tropical conditions were close to those obtained pollination depends on flowering timing. A difference of 04
in Romania on improved maize varieties by Viorica et al. to 05 days of flowering time between the pollen source and
(2017) who noted a xenia percentage of 44.9% at 1 m from the recipient reduces the pollen flow pear to 50%. It is clear
pollen source and 0.33% at 20 m. However, Bannert and that flowering synchronization between neighboring fields
Stamp (2007) investigated the effectiveness of distance in is the main factor influencing cross-pollination (Messenger
preventing out-crossing in maize and showed that the rate et al., 2006). The size and the ratio between the pollen
of cross-pollination ranged from 3% to 15% at 0.8 m from source and the receiving field also influence the level of
the donor. There is a direct correlation between the level of cross-pollination. The deeper the receiving field, the lower
cross-pollination and to distance to source (R=0.928). At the level of cross-pollination of the crop production (Ma et
larger distance from the source, Aylor et al. (2003) noted a al., 2004). Cross-pollination is considered to be responsible
cross-pollination rate of 0.1 % at 50 m. Raynor et al. (1972) for much of the gene flow in maize (Devos et al., 2007).
estimated that less than 1% of maize pollen grains traveled Gene flow influences reproductive success and fitness of
beyond 60 m, considering that maize pollen is the largest individuals, and determines the genetic structure of the
and heaviest of the Poaceae pollinated species. Dispersal population.

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Table 3. Analysis of variance for cross-pollination rate within the recipient field of maize tested for eight distances from
pollen-donor source and four wind directions
Source of variation Df SS MS F-value
Blocks 2 13.57 6.78 0.78ns
Distance from source (D) 7 2668.32 381.18 43.86***
Wind direction (Wd) 3 483.90 161.30 18.56***
Interaction D × Wd 21 181.22 8.63 0.99ns
Residual 62 539.04 8.69
Df: Degree of freedom; SS: Sum of square; MS: Mean of square; ns: not significant at 5%; ***: indicates significance at 0.1%.

Table 4. Impact of distance from pollen-source and wind direction on the cross-pollination rate (%) within four recipient
maize synthetic varieties
Gap from pollen Percentage of outcrossing (%)
source (recipient North South East West Average for
variety) distance
1.5 m (CMS 9015) 44.64±0.28 34.67±0.41 34.94±0.38 40.24±0.31 38.62±0.34a
2.5 m (CMS 9015) 33.33±1.51 23.04±0.62 25.24±1.05 28.23±0.99 27.46±1.04b
4.0 m (CMS 8501) 25.31±0.53 14.88±0.57 15.32±0.60 21.92±0.38 19.35±0.52c
5.0 m (CMS 8501) 14.97±0.59 11.99±0.85 12.47±0.85 13.35±0.44 13.19±0.68d
6.5 m (CMS 2019) 11.92±0.85 9.13±0.39 8.93±0.16 11.02±0.62 10.25±0.50e
7.5m (CMS 2019) 9.11±0.41 7.15±0.56 6.63±0.65 7.95±0.55 7.71±0.54f
9.0 m (Shaba) 8.21±0.41 5.39±0.56 5.95±0.65 7.59±0.55 6.78±0.28fg
10.0 m (Shaba) 6.72±0.43 4.03±0.37 4.65±0.25 5.81±1.00 5.30±0.51g
Average value for 19.27±0.61A 13.78±0.51C 14.26±0.50C 17.01±0.58B 16.08±0.55
direction
Means with the same subscript within the same column or line do not differ significantly at 5%.

45
40
35
Cross-pollination rate

R= 0.928
30
25
20
pollution
15
10
5
0
1 2 3 4 5 6 7 8 9 10 11 12 13
Distance from pollen source

Fig. 2: Impact of gap from pollen source on the cross-pollination rate of recipient varieties.

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25

a
20
Cross pollinatio rate

b
c c
15

10

0
NORTH SOUTH EAST WEST

Directions

Fig. 3: Impact of wind direction on cross-pollination rate of recipient maize varieties.

IV. CONCLUSION [7] Della P.G., Ederle D., Bucchini L., Prandi. M., Verderio A.
& Pozzi C., 2008. Maize pollen mediated gene flow in the Po
The main focus of this study was to gather
valley (Italy): source-recipient distance and effect of flowering
information’s about adequate separation distances to ensure time. European. Journal. Agronomy, 28: 255-265.
coexistence and about the dependence of cross-pollination [8] Devos Y., Reheul D. & De Schrijver A., 2005. The co-
on the distance within the maize field. Results obtained existence between transgenic and nontransgenic maize in the
showed the level of genetic pollution is highest near the European Union: a focus on pollen flow and cross-
pollen source and in the northern direction. At the distance fertilization. In: Environmental Biosafety Research, 4: 71-87.
of 10 m from the pollen source, the average level of genetic [9] Devos Y., Reheul D., Thas O., De Clereq E.M., Cougnon
pollution decreased significantly in the South direction. We M. & Cordeman S.K., 2007. Implementing isolation
perimeters around genetically modified maize fields.
can recommend the cultivation at a distance of more than 10
Agronomy. Sustain Dev., 27: 155-165.
m between maize fields to secure the coexistence of
[10] Djoufack M.V., Fontaine B., Martiny N. & Tsalefac M.,
genotypes. 2012. Climatic and demographic determinants of vegetation
cover in Northen Cameroon. International Journal of Remote
Sensing, 6904-6926.
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