Research Article | | Peer-Reviewed

Interactive Effects of Substrate and Length on the Ability of Root Segment Cuttings of Burkea Africana (Hook) and Uvaria Chamae (P. Beauv) to Regenerate

Received: 6 February 2026     Accepted: 20 February 2026     Published: 10 March 2026
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Abstract

The Guinean Savanah Highlands of Adamawa is replete with multipurpose tree species, among which Burkea africana and Uvaria chamae are particularly noteworthy. Despite their importance, they remain in the wild and are subjected to overexploitation. The present study aims to contribute to the domestication of these species by root segment propagation. Specifically, the study aims to evaluate the seasonal variations in carbohydrate reserves (starch, soluble sugars, and sucrose) in other to determine the most favorable period for root cutting collection, assess the effect of substrate and length of root segment cuttings on the budding and rooting capacity of these species, evaluate the effect of carbohydrate content on bud emergence, root formation, callus induction, and control response in cuttings. For the seasonal fluctuation of carbohydrate, the experimental design was a complete randomized design with one factor represented by the season, and two replications. In the case the root propagation, the experimental design was a split-plot with three replications. The main treatment comprised three substrates (sand/sawdust, black soil/sawdust, black soil), while the sub-treatments were represented by three lengths of root segments cuttings (RSC) (10, 15, 20cm). The experimental unit consisted of 10 cuttings. Results showed that the onset of the rainy season coincides with peak of starch, soluble sugars and sucrose for Burkea africana and Uvaria chamae, marking the most favorable period for root cutting collection. The budding rate of Burkea africana showed significant variation (0.04 < 0.05), with values of 2.22 ± 1.96% in a black soil/sawdust mixture and 14.44 ± 12.36% in black soil after 26 weeks. For Uvaria chamae the best substrate was the mixture of black soil/sawdust (63.33 ± 45.27%). The number of leaves for Burkea africana was substantially higher in black soil (4.27 ± 2.16). The difference was statistically significant (0.008 < 0.01). For Burkea africana and Uvaria chamae, the optimal cutting length for budding was 20 cm (11.11 ± 10.52%, 81.11 ± 26.20%). The rooting rate of Uvaria chamae exhibited considerable variability the best rate was those of the black soil/sawdust substrate (22.22 ± 20.33%). The rooting rate varied from 5.55 ± 4.26% for cuttings of 10 cm to 23.33 ± 21.79% for those of 20 cm.Budded cuttings clustered with soluble sugars and sucrose. Rooted cuttings correlate negatively with all carbohydrate. Control cuttings were closely associated with starch. All these informations are important to develop scales and strategies toward the domestication of this species.

Published in American Journal of Agriculture and Forestry (Volume 14, Issue 2)
DOI 10.11648/j.ajaf.20261402.11
Page(s) 74-91
Creative Commons

This is an Open Access article, distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution and reproduction in any medium or format, provided the original work is properly cited.

Copyright

Copyright © The Author(s), 2026. Published by Science Publishing Group

Keywords

Burkea Africana, Uvaria Chamae, Guinean Savannah Highlands, Vegetative Propagation, RSC

1. Introduction
Burkea africana, commonly known as Kokobi in Fulani, belongs to the Fabaceae family and is a widespread species across tropical Africa. Its wood is hard and heavy, making it highly suitable for various construction purposes such as bridges, railway sleepers, furniture, fences, and tool handles. It is also extensively used as firewood and for charcoal production . Notably, the heartwood is resistant to fungal decay . Beyond its structural uses, the bark, roots, and leaves of B. africana are widely used in traditional medicine . Uvaria chamae, known as Banane wandou in Fulani, belongs to the Annonaceae family. Its ripe yellow fruits contain a sweet, palatable pulp that is commonly consumed. Additionally, both the leaves and roots are used in ethnomedicine to treat malaria, typhoid and diabetes, highlighting the species' broad therapeutic potential . These species however, are increasingly threatened by anthropogenic pressures, which are accelerating the depletion of both timber and non-timber forest products . Therefore, it is crucial to preserve local species of socio-economic importance not only in forest reserves but also within agropastoral systems near the communities that rely on them . To alleviate pressure on natural resources, local communities in developing countries have adopted domestication and agroforestry practices as immediate and cost-effective strategies. Sexual reproduction remains the primary propagation technique for timber species and biodiversity enhancement (Moupela et al., 2013; Meunier et al., 2008). However, germination and seedling development are often poor in the dry tropical savannah . Consequently, seed multiplication remains unreliable for many rural communities in Africa due to limited seed availability, financial constraints, and predation by ants, birds, and other animals . Vegetative propagation offers a faster and more affordable alternative . It is a promising adaptive strategy for these species, allowing them to withstand environmental disturbances and climatic variability . By capturing germplasm from wild populations, vegetative propagation supports the domestication, conservation, and sustained availability of these plants for local communities. It replicates the genetic traits of the parent plant and provides several advantages, including rapid plant production, early fruiting, and smaller plant size .
The main objective of this study is to contribute to the domestication of this species in order to protect and introduce them in existing farmer's production systems. Specifically:
1) optimal sampling period for root segment cuttings collection;
2) evaluate the effect of substrate on the budding and rooting ability;
3) assess the effect of cutting length on the budding and rooting ability;
4) effect of Carbohydrate content on developmental stages of cuttings.
2. Materials and Methods
2.1. Study Site Description
The research was carried out in the Guinean Savannah highland of Adamawa, located between latitude 6° and 8° N and longitude 11° 30' and 15° 30' E. The region is defined by two major ecological limits: the Sudanian savannah to the north and the semi-deciduous Guinean forest to the south. Vegetation is predominantly shrub and tree savannah, largely dominated by Daniellia oliveri and Lophira lanceolata . The disruption of the vegetation is significantly influenced by human activities. The region is characterised by a guinean climate, composed of two distinct seasons: a rainy season from April to October and a dry season from November to March . The area is home to numerous ethnic groups, including the Fulani, Mboums, Pères, Koutines, Haoussas, Niza’as, and Dourous . The soil is characterized by a red ferritic structure developed on old basalt .
2.2. Description of the Nursery and the Non-Mist Propagator
The root segment cuttings experimentation was conducted at the nursery of the laboratory of Ecology and Sustainable Development of the University of Ngaoundere. The cuttings were propagated in a non-mist propagator placed beneath a sheet shed, which provides shade. Six transparent sheet sheds are positioned within the roof to filter the light. Each non-mist propagator is constructed from local materials and is shaped like a parallelepiped, subdivided into three compartments. The frame is made of wood and covered with transparent polyethylene, which ensures favorable conditions for the development of cuttings. The relative humidity in the non-mist propagators is between 80 and 100%, while the temperature varies from 23 to 28°C . The internal configuration of the non-mist propagators comprises the following layers, from the bottom to the top: a thin layer of fine sand, large pebbles, medium pebbles, gravel, sand and finally rooting substrates . A PVC pipe is fixed to the corner of the non-mist propagators to facilitate regular gauging of the water level.
2.3. Methodology
2.3.1. Optimal Sampling Period for Root Segment Cuttings Collection
The seasonal fluctuation of carbohydrates is key to identifying the optimal sampling period for root segment cuttings in vegetative propagation. However, this remains underdocumented in the Guinean savannah highlands of Adamawa, especially for, Burkea africana, and Uvaria chamae. To fill this gap, a year-long root sampling program was carried out to monitor seasonal carbohydrate variation in these species.
Monthly root samples (0.5 kg) were collected from 15 mature individuals per species, with U. chamae sampled at Nyambaka and B. africana at Mberem. Sampling followed the region’s climatic calendar: early dry (Oct–Dec), late dry (Jan–Mar), early rainy (Apr–Jun), and late rainy (Jul–Sep).
Samples were air-dried for a month, ground into powder, and stored in hermetically sealed, labeled bags indicating species, date, and location. Biochemical analyses were conducted at the University of Maroua’s Laboratory of Biochemistry and Organic Chemistry to quantify key carbohydrates linked to bud initiation and rooting potential. This helped identify the optimal season for propagation via root cuttings. Three main sugars were chosen in this operation.one non-stuctural sugar represented by starch and two structural sugars (soluble sugars and sucrose).
(i). Determination of Soluble Sugar Content
Principle:
Based on , reducing sugars (glucose, galactose, fructose) form furfural under heat, which reacts with phenolic compounds (ADNS) to produce a red-brown complex measurable at 530 nm.
Extraction:
100 mg of root powder was mixed with 5 mL of 80% ethanol in a 10 mL tube, stirred for 24 h, and filtered through Whatman No. 4 paper.
Procedure:
In a Pyrex tube, 0.5 mL of sample or glucose standard (10 mg/mL) and 1 mL of ADNS reagent were mixed, heated in boiling water (90 °C, 5 min), then cooled and diluted with 1 mL distilled water. Tests were done in triplicate. A galactose-based calibration curve (0–500 µg/mL) was used to quantify sugars, expressed in mg galactose equivalent per 100 g dry matter (mgEG/100g DM).
SSR (gEG100gDM)=DO*fd*VPe*a
where SSR = Content of soluble reducing sugars; DO = Optical density; fd = Dilution factor; V = Total extraction volume; Pe = Sample intake; a = Slope of the calibration curve
(ii). Determination of Sucrose Content
Using protocol, the ethanol filtrate was hydrolyzed with 0.5 mL of 1M HCl, heated at 90 °C for 10 min, then neutralized with 0.6 mL of 1M NaOH. After cooling, the same reducing sugar assay was applied. Sucrose was estimated as:
Sucrose=Total soluble sugars − Reducing sugars
(iii). Determination of Resistant Starch Content
Principle:
Based on protocol, iodine binds to amylose (blue complex, λmax = 630 nm) and amylopectin (brown, λmax = 548 nm). Absorbance at 580 nm reflects total starch content.
Protocol:
0.1 g of root powder was treated with 3 mL of 1N KOH, homogenized (10 min), neutralized with 3 mL of 1N HCl, boiled (90 °C, 15 min), cooled, and diluted to 4 mL. After filtration and 10× dilution, 0.5 mL of each sample or starch standard was mixed with 0.5 mL water and 1 mL I₂/KI solution. Tubes were incubated at room temperature for 10 min. A starch calibration curve (0–400 µg/mL) was used to determine total starch content, expressed in g per 100 g of material.
2.3.2. Root Segment Propagation
The methodological approach involved the partial and carefull excavation of the root systems of 15 adult trees in the first hour of the morning. The excavation process was conducted in Nyambaka. Following the excavation process, the cuttings segments were immediately placed in a cooler with ice blocks and transported from the field to the nursery to minimize water loss . Upon arrival at the nursery of the Laboratory of Ecology and Sustainable Development of the University of Ngaoundere, the cuttings segments were trimmed to three distinct lengths: 10, 15, and 20 cm (Figure 1) and marked at the proximal end, then inserted vertically in the non-mist propagator. The different substrates had already been prepared and were of three kinds (sand/sawdust, black soil/sawdust and black soil).
Figure 1. Different length of cuttings.
For the mixture of sand and sawdust and the mixture of black soil and sawdust, the substrates were prepared in a 50:50%. Approximately 1 cm of the proximal end of the cuttings was left exposed beyond the substrate. The sawdust was obtained from a local sawmill and allowed to decompose. The black soil was obtained from the nursery, while the sand was collected from the river in the vicinity of Ndom, The experimental design was a split plot with three replications, with the main treatment represented by the substrate and the secondary, the length of root segments cuttings. The experimental unit was composed of 10 cuttings.
2.3.3. Carbohydrate Content on Developmental Stages of Cuttings
During the root cutting propagation process, three types of cuttings: control cuttings (cuttings which did not produce adventitious shoots), callus cuttings, budded cuttings, and rooted cuttings. Samples were air-dried for one month, ground into fine powder, and stored in hermetically sealed, labeled bags indicating species, date, and location. Biochemical analyses were carried out at the University of Maroua’s Laboratory of Biochemistry and Organic Chemistry to quantify key carbohydrates (starch, soluble sugars, and sucrose). The experimental design followed a completely randomized design with one factor and two repetitions.
2.3.4. Data Collection and Analysis
Data were collected on a weekly basis in order to monitor the budding process (from the date of appearance of the first bud) and at the end of the experiment in order to assess the rooting success. The data set comprised the following variables: the number of cuttings that exhibited buds, the number of leafy axes, the height of the leafy shoots, the number of leaves per shoot, the number of rooted cuttings, the number of roots per cutting, and the length of adventitious roots. Furthermore, the aforementioned growth parameters were evaluated at the end of the experimental period. A root segment cutting (RSC) is deemed to have successfully rooted if the length of the root is equal to or greater than 1 cm. In the event that this criterion is not met, the cutting is returned to the substrate . The data collected were subjected to an analysis of variance. Significant means were separated using the Duncan multiple range test. The statistical software used for the analysis of variance was Statgraphics Plus 5.0. Excel was used to generate graphs.
3. Results
3.1. Seasonal Changes in Carbohydrate Content in the Guinean Savannah Highlands
3.1.1. Starch Content
The starch content of Burkea africana exhibited a significant seasonal variation (0.001 < 0.01), with values ranging from 4.33 ± 0.94 g/100 g DM at the end of the rainy season to 6.39 ± 0.84 g/100 g DM at the beginning of the rainy season (Table 1a). Similarly, Uvaria chamae showed a significant increase in starch content (0.000 < 0.001). Ranging from 4.57 ± 0.49 g/100 g DM at the end of the rainy season to 8.49 ± 0.73 g/100 g DM at the beginning of the rainy season (Table 1b).
a) Burkea africana

Seasons

Dry season

Rainy season

P-value

Beginning

End

Beginning

End

Starch (g/100 g DM)

5.65 ± 0.98bc

4.63±0.59ab

6.39±0.84c

4.33±0.94a

0.001

b) Uvaria chamae

Seasons

Dry season

Rainy season

P-value

Beginning

End

Beginning

End

Starch (g/100 g DM)

4.90 ± 0.64a

7.32±0.87b

8.49±0.73c

4.57±0.49a

0.0000

Means with identical letter do not differ significantly at P ≤ 0.05 according to Duncan’s Multiple Range Test
3.1.2. Soluble Sugar Content
The soluble sugar content for Burkea africana showed values ranging from 2.23 ± 0.23 g/100 g DM at the end of the rainy season to 4.16 ± 0.91 g/100 g DM at the beginning of the rainy season (Table 2a). The analysis of variance indicates a significant seasonal difference (0.000 < 0.001). Uvaria chamae demonstrated also a marked seasonal effect on soluble sugar content, varying from 2.75 ± 0.21 g/100 g DM at the onset of the rainy season to a peak of 6.41 ± 0.60 g/100 g DM at the conclusion of the rainy season, and this variation was highly significant (0.000 < 0.001) (Table 2b).
a) Burkea africana

Seasons

Dry season

Rainy season

P-value

Beginning

End

Beginning

End

Soluble sugars (g/100 g DM)

2.55 ± 0.31ab

2.97±0.51c

4.16±0.91d

2.23±0.23a

0.0000

b) Uvaria chamae

Seasons

Dry season

Rainy season

P-value

Beginning

End

Beginning

End

Soluble sugars (g/100 g DM)

3.72 ± 0.41b

5.62±0.76c

6.41±0.60d

2.75±0.21a

0.0000

Means with identical letter do not differ significantly at P ≤ 0.05 according to Duncan’s Multiple Range Test
3.1.3. Sucrose Content
The sucrose content of Burkea africana exhibited a broader range of sucrose content, varying from 1.07 ± 0.20 g/100 g DM at the end of the rainy season to 2.12 ± 0.48 g/100 g DM at the beginning of the subsequent rainy season (Table 3a). The differences observed between seasons were statistically significant (0.001< 0.01), indicating that seasonal factors strongly influence sucrose accumulation in this species. Similarly, Uvaria chamae showed pronounced seasonal fluctuations in sucrose content, with values ranging from 1.50 ± 0.27 g/100 g DM at the onset of the rainy season to a peak of 4.48 ± 1.06 g/100 g DM by its end (Table 3b). The analysis of variance indicates a significant variation (0.001 < 0.01).
a) Amblygonocarpus andongensis

Seasons

Dry season

Rainy season

P-value

Beginning

End

Beginning

End

Sucrose (g/100 g DM)

1.34 ± 0.12a

1.83±0.16b

2.12±0.48b

1.07±0.20a

0.0000

b) Uvaria chamae

Seasons

Dry season

Rainy season

P-value

Beginning

End

Beginning

End

Sucrose (g/100 g DM)

2.78 ± 0.05b

3.33±0.78c

4.48±1.06d

1.50±0.27a

0.0000

Means with identical letter do not differ significantly at P ≤ 0.05 according to Duncan’s Multiple Range Test
3.2. Budding Rate
The sprouting time for these species is respectevely 2 and 12 weeks, for Burkea africa and Uvaria chamae which is the period between the placement of the cutting into cultivation and the initial appearance of the bud (Figure 2).
Figure 2. Budded cuttings.
3.2.1. Effect of Substrate
The budding rate varies from 2.22 ± 1.96% in a black soil/sawdust mixture to 14.44 ± 12.36% in black soil alone after 26 weeks (Figure 3a). The budding rate of Burkea africana showed significant variation (0.04 < 0.05). In contrast, Uvaria chamae exhibited relatively stable budding rates after 17 weeks, with non-significant fluctuations (0.90 > 0.05) from 53.33 ± 50.74% in black soil to 63.33 ± 45.27% in the black soil/sawdust mixture (Figure 3b).
Figure 3. Effect of substrate on the budding rate.
3.2.2. Effect of length
Variation in budding rates across cutting lengths was observed in both Burkea africana and Uvaria chamae, though these differences were not statistically significant respectevely (0.25 > 0.05 and 0.19 > 0.05). In B. africana, budding success exhibited a modest increase from 4.44 ± 2.69% in 10 cm cuttings to 11.11 ± 10.52% in 20 cm cuttings (Figure 4a). In contrast, U. chamae showed a more pronounced increase in budding rates, rising from 36.66 ± 34.47% to 81.11 ± 26.20% over the same cutting length range (Figure 4b)
Figure 4. Effect of length of the cuttings on the budding rate.
3.2.3. Effect of the Interaction Substrate Length
For Burkea africana, the success rates ranged from 0 ± 0% for the 10 cm and 20 cm cuttings placed in the black soil/sawdust mixture, up to 26.66 ± 15.27% for the 20 cm cuttings inserted in black soil alone (Figure 5a). The analysis of variance indicated that these differences were not statistically significant (0.13 > 0.05). For Uvaria chamae, the results showed a wider range of values. The 10 cm cuttings inserted in sand/sawdust and black soil substrates had an average success rate of 33.33 ± 27.73%, while the 20 cm cuttings inserted in the sand/sawdust substrate achieved a much higher mean of 96.66 ± 5.77% (Figure 5b). However, the analysis of variance revealed no statistically significant differences (0.96 > 0.05).
Figure 5. Budding rate following the interaction substrate*length of RSC.
3.3. Budding Parameters
3.3.1. Effect of Substrate
(i). Number of Shoots
For Burkea africana, the number of leafy shoots was significantly influenced by the substrate (0.02 < 0.05). Shoot numbers ranged from 0.52 ± 0.40 in the black soil/sawdust mixture to 2.29 ± 1.43 in pure black soil (Table 4a). In contrast, for Uvaria chamae, although the number of leafy shoots varied from 17.66 ± 15.46 in black soil to 23.44 ± 19.05 in sand/sawdust (Table 4b). The difference was not statistically significant (0.79 > 0.05).
(ii). Height of Shoots
B. africana exhibited relatively low and less variable shoot heights, ranging from 1.35 ± 1.10 cm in black soil/sawdust to 2.49 ± 1.25 cm in black soil (Table 4a). The variance analysis did not reveal significant differences among substrates (0.59 > 0.05). Similarly, U. chamae showed fluctuations in shoot height, with means of 5.63 ± 5.45 cm in black soil and 8.29 ± 5.68 cm in black soil/sawdust (Table 4b). However, the difference was not statistically significant (0.62 > 0.05).
(iii). Number of Leaves
The number of leaves exhibited notable variation among treatments, with Burkea africana showing a significant response to substrate type. Specifically, the mean leaf count was substantially higher in black soil (4.27 ± 2.16) compared to the black soil–sawdust mixture (0.91 ± 0.58) (Table 4a). The difference was statistically significant (0.008 < 0.01), indicating that the black soil alone provided more favorable conditions for leaf development in this species. For Uvaria chamae, leaf number varied widely, with a mean of 70.33 ± 69.97 in black soil and 92.11 ± 89.45 in the black soil/sawdust substrate (Table 4b). Despite this apparent increase, the high standard deviations and overlapping confidence intervals resulted in no statistically significant difference between treatments (0.81 > 0.05).
a) Burkea africana

Substrate

Number of shoots

Height of shoots

Number of leaves

Sand/Sawdust

1.46 ± 1.40b

1.93 ± 1.66a

2.27 ± 2.27b

Black soil

2.29 ± 1.43b

2.49 ± 1.25a

4.27 ± 2.16c

Black soil/Sawdust

0.52 ± 0.40a

1.35 ± 1.10a

0.91 ± 0.58a

P-value

0.02

0.59

0.007

b) Uvaria chamae

Substrate

Number of shoots

Height of shoots

Number of leaves

Sand/Sawdust

23.44 ± 19.05a

7.23 ± 6.12a

76.11 ± 61.64a

Black soil

17.66 ± 17.67a

5.63 ± 5.45a

70.33 ± 69.97a

Black soil/Sawdust

18.88 ± 16.98a

8.29 ± 5.68a

92.11 ± 89.45a

P-value

0.31

0.48

0.81

Means with identical letter do not differ significantly at P ≤ 0.05 according to Duncan’s Multiple Range Test
3.3.2. Effect of Length
(i). Number of Shoots
For the number of leafy shoots for Burkea africana, the variation in leafy shoot production between 10 cm (0.81 ± 0.46) and 15 cm (1.81 ± 1.13) cuttings was not statistically significant (0.22 > 0.05) (Table 5a). In Uvaria chamae, the number of leafy shoots varied substantially from 8.33 ± 6.46 in 10 cm cuttings to 31.44 ± 17.57 in 20 cm cuttings (Table 5b). The analysis of variance indicated a significant difference (0.02 < 0.05).
(ii). Height of Shoots
The height of shoot of Burkea africana, demonstrated a fluctuating pattern, ranging from 0.65 ± 0.27 cm in 10 cm cuttings to 3.24 ± 2.84 cm for 15 cm cuttings (Table 5a). However, analysis of variance indicated no significant difference among the treatments (0.08 > 0.05). Similarly, in Uvaria chamae, shoot height varied from 4.03 ± 2.27 cm in 10 cm cuttings to 10.00 ± 4.72 cm in 20 cm cuttings (Table 5b), yet the differences were not statistically significant (0.12 > 0.05).
(iii). Number of Leaves
The number of leaves in Burkea africana displayed a different pattern, with leaf counts ranging from 1.33 ± 1.08 in 10 cm cuttings to 3.25 ± 2.07 in 20 cm cuttings (Table 5a); however, the differences were not statistically significant (0.12 > 0.05). For Uvaria chamae, the number of leaves varied significantly with cutting length, increasing from 26.22 ± 16.58 for 10 cm cuttings to 121.33 ± 78.68 in 20 cm cuttings (Table 5b), with the effect confirmed by analysis of variance (0.03 < 0.05).
a) Burkea africana

Length of RSC (cm)

Number of shoots

Height of shoots (cm)

Number of leaves

10

0.81 ± 0.46a

0.65 ± 0.27a

1.33 ± 1.08a

15

1.81 ± 1.13a

3.24 ± 2.84a

3.25 ± 2.07a

20

1.66 ± 1.52a

1.88 ± 1.37a

2.86 ± 2.59a

P-value

0.72

0.70

0.69

b) Uvaria chamae

Length of RSC (cm)

Number of shoots

Height of shoots(cm)

Number of leaves

10

8.33 ± 6.46a

4.03 ± 2.27a

26.22 ± 16.58a

15

20.22 ± 15.57ab

7.12 ± 5.85a

91 ± 72.48b

20

31.44 ± 17.57b

10.00 ± 4.72a

121.33 ± 78.68c

P-value

0.02

0.12

0.03

Means with identical letter do not differ significantly at P ≤ 0.05 according to Duncan’s Multiple Range Test
3.4. Rooting Rate
3.4.1. Effect of Substrate
The rooting rate of Uvaria chamae exhibited considerable variability, with values ranging from 11.11 ± 10.64% in the black soil to 22.22 ± 20.33% in the black soil/sawdust substrate (Table 6). The analysis of variance did not indicate significant variation (0.44 > 0.05). In the case of Burkea africana after 24 weeks none of the cutting profuse adventitious roots.
Table 6. Effect of substrate on the rooting rate.

Substrates

Sand/Sawdust

Black soil

Black soil/Sawdust

Mean

14.44 ± 14.24a

11.11 ± 10.64a

22.22 ± 20.33a

P-value

0.44

Means with identical letter do not differ significantly at P ≤ 0.05 according to Duncan’s Multiple Range Test
3.4.2. Effect of Length of Root Segments Cuttings
For Uvaria chamae, the rooting rate varied from 5.55 ± 4.26% for cuttings of 10 cm to 23.33 ± 21.79% for those of 20 cm (Table 7). However, the analysis of variance did not indicate significant variation (0.13 > 0.05).
Table 7. Effect of length of cutting on the rooting rate.

Length of RSC (cm)

10

15

20

Mean

5.55 ± 4.26a

18.88 ± 16.04a

23.33 ± 21.79a

P-value

0.13

Means with identical letter do not differ significantly at P ≤ 0.05 according to Duncan’s Multiple Range Test
Figure 6. Rooted cutting of Uvaria chamae.
3.4.3. Effect of Substrate Length Interaction
For U. chamae, the rooting rate exhibited fluctuations, ranging from 3.33 ± 1.77% for the cutting of 10 cm inserted in the sand/sawdust and black soil to 26.66 ± 5.77% for those of 20 cm put into the sand/sawdust mixture (Fig7). The analysis of variance did not indicate a significant variatin (0.48 > 0.05).
Figure 7. Rooting percentage according the interaction substrate*length.
3.5. Rooting Parameters
3.5.1. Effect of Substrate
(i). Number of Roots
In Uvaria chamae, the number of roots varied from 1.55 ± 0.9 in cuttings placed in black soil to 2.88 ± 2.75 in those placed in the sand/sawdust mixture (Table 8). However, analysis of variance revealed no significant difference between the treatments (0.9 > 0.05).
(ii). Length of Roots
The length of root of U. chamae, exhibited fluctuations, with the cutting inserted in the black soil displaying a value of 1.74 ± 1.22 cm, while those placed in the sand/sawdust mixture recorded a value of 5.56 ± 3.07 cm (Table 8). The analysis of variance did not indicate a significant variation (0.43 > 0.05).
Table 8. Effect of substrate on rooting parameters.

Substrates

Number of roots

Length of roots

Sand/Sawdust

2.88 ± 2.75a

5.56 ± 3.07a

Black soil

1.55 ± 0.9a

1.74 ± 1.22a

Blacksoil/Sawdust

2.77 ± 2.38a

3.03 ± 2.54a

P-value

0.50

0.43

Means with identical letter do not differ significantly at P ≤ 0.05 according to Duncan’s Multiple Range Test
3.5.2. Effect of Length of Cuttings
(i). Number of Roots
The number of roots of Uvaria chamae varied, with values ranging from 1 ± 0.22 for the 10 cm cuttings to 2.88 ± 2.42 for those of 20 cm (Table 9). The analysis of variance indicated no significant variation (0.16 > 0.05).
(ii). Length of Roots
The length of roots of U. chamae, demonstrated fluctuations, with values ranging from 1.67 ± 1.15 cm for the 10 cm cuttings to 5.31 ± 3.09 cm for those of 15 cm (Table 9). The analysis of variance indicated that no significant variation was present (0.12 > 0.05).
Table 9. Effect of substrate on rooting parameters.

Lenght of RSC (cm)

Number of roots

Length of roots (cm)

10

1 ± 0.22a

1.67 ± 1.15a

15

3.33 ± 2.5a

5.31 ± 3.09a

20

2.88 ± 2.42a

3.33 ± 2.72a

P-value

0.16

0.12

Means with identical letter do not differ significantly at P ≤ 0.05 according to Duncan’s Multiple Range Test
3.5.3. Carbohydrate content Dynamics on Developmental Stages of Cuttings
Following the propagation stages, some relationships are developped between carbohydrates and cutting stages and among carbohydrates. The correlation analysis revealed for Uvaria chamae strong positive relationships among starch, soluble sugars, and sucrose (r = 0.662, 0,864, –0.949), indicating that these carbohydrate fractions are metabolically interconnected and tend to accumulate together (Table 7). In contrast, the budding and rooting rates were generally negatively correlated with carbohydrate content. Rooting rate showed particularly strong negative associations with starch (r = –0.947), soluble sugars (r = –0.867), and sucrose (r = –0.980). Budding rate exhibited a moderate negative correlation with starch (r = –0.751) and a weak negative correlation with sucrose (r = –0.317) (Table 10). The moderate positive correlation between budding and rooting rates (r = 0.500) further indicates that tissues capable of initiating buds efficiently may also exhibit enhanced rooting potential, likely. Collectively, the results imply that while carbohydrates are essential for growth, their utilization rather than absolute abundance is critical for successful bud and root formation.
Table 10. Relationships between carbohydrates and developmental cuttings stages of Uvaria chamae.

Variables

Starch

Soluble sugars

Sucrose

Budding rate

Rooting rate

Starch

1

Soluble sugars

0,662

1

Sucrose

0,864

0,949

1

Budding rate

-0,751

-0,002

-0,317

1

Rooting rate

-0,947

-0,867

-0,980

0,500

1

The correlation analysis in Burkea africana revealed perfect relationships among starch, soluble sugars, sucrose, and budding rate, with coefficients of ±1 (r = ±1) (Table 11), indicating tightly coupled metabolic associations. Starch and sucrose were perfectly positively correlated, while both were perfectly negatively correlated with soluble sugars. Budding rate exhibited a perfect positive correlation with soluble sugars and perfect negative correlations with starch and sucrose.
Table 11. Relationships between carbohydrates and developmental cuttings stages of Burkea Africana.

Variables

Starch

Soluble sugars

Sucrose

Budding rate

Starch

1

Soluble sugars

-1,000

1

Sucrose

1,000

-1,000

1

Budding rate

-1,000

1,000

-1,000

1

Principal component analysis (PCA) gives supplementary explanations regarding these relationships between carbohydrate fractions (starch, soluble sugars, and sucrose) and the developmental stages of cuttings in Uvaria chamae and Burkea africana. For U. chamae, axes F1 and F2 together explained 100% of the total variance, with F1 accounting for 88.53% and F2 for 11.47%. Soluble sugars and sucrose were strongly correlated with F1 and moderately with F2, while starch exhibited a positive correlation with F1 and a negative correlation with F2. Budded cuttings clustered near the vectors for soluble sugars and sucrose, while rooted cuttings were positioned away from all carbohydrate vectors. Control cuttings (that did not emit adventitious shoot) were closely associated with starch (Figure 8a). For B. africana, the PCA revealed a one-dimensional distribution, with 100% of the variance explained by F1 alone. All carbohydrate fractions aligned along this single axis. Budded cuttings were located on the negative side of F1, near soluble sugars, whereas control cuttings were positioned on the positive side, closer to starch and sucrose. Rooted and callus cuttings were not represented in the dataset (Figure 8b).
Figure 8. Principal Component Analysis (PCA) of Carbohydrate content and physiological state of different types of cuttings.
4. Discussion
4.1. Optimal Sampling Period for Root Segment Cuttings Collection
The discrepancy in seasonal peaks of soluble sugar concentrations may be attributed to species-specific physiological responses and environmental conditions. This may explain the divergence from the findings of , who reported the highest soluble sugars levels in Bombax costatum at the onset of the dry season. Conversely, the current results agree with those of , who observed peak soluble sugar accumulation during the rainy season, corresponding to the autumn period, in Populus tremuloides. The seasonal dynamics of soluble sugar content likely reflect plant responses to water availability. During dry periods, water stress typically limits photosynthetic activity, yet it often leads to increased concentrations of soluble sugars in plant tissues. These sugars function as osmolytes, helping cells maintain turgor pressure under drought conditions. Moreover, plants may hydrolyze stored starch reserves into soluble sugars to counteract the osmotic stress induced by water deficit . Conversely, during the rainy season, enhanced water availability promotes higher rates of photosynthesis and carbohydrate synthesis. The surplus sugars generated under these favorable conditions are utilized to support growth processes such as leaf flushing.
In their natural environment, Burkea africana and Uvaria chamae exhibit marked phenological cycles that correspond to the distinct raining and dry seasons of the region. The raining season, which extends from May to October, is characterized by high rainfall and moderate temperatures, while the dry season, from November to April, is marked by little or no rain and higher evapotranspiration. During the early wet season (May–June), both species initiate active vegetative growth, with leaf flushing and shoot elongation occurring soon after the first rains . This is followed by flowering and fruit development in the mid- to late wet season (July–October). In contrast, during the dry season (November–April), the phenological activity declines: leaves senesce and are shed, reproductive activity ceases, and the plants enter a state of dormancy to withstand moisture stress. According to , Burkea africana begins leaf flushing shortly after the onset of the rains in May, flowers later in the wet season, and sheds its leaves with the arrival of the dry season. Similarly, Uvaria chamae, as reported by , produces new shoots and flowers predominantly during the rainy months, with reduced growth and partial dormancy during the dry period.
The patterns of sucrose accumulation in these species follow the seasonal phenological changes and are consistent with the findings of in Populus tremuloides, where seasonal variability in moisture availability strongly influenced carbohydrate dynamics. In both Burkea africana and Uvaria chamae, the onset of the wet season and increased photosynthesis promote sucrose synthesis in the leaves, which is then translocated to developing tissues to support growth, reproduction, and storage during the growing season.
4.2. Root Segment Propagation
4.2.1. Budding
Burkea africana began to sprout after 12 weeks. In contrast, Uvaria chamae exhibited rapid bud emergence within just 2 weeks. These findings underscore the significant interspecific differences in vegetative regeneration dynamics and suggest a hierarchy of physiological responsiveness among the species evaluated. The rapid sprouting observed in Uvaria chamae are nearer to those observed by , who reported mean sprouting periods of three weeks for Bombax costatum and Diospyros mespiliformis, respectively. This precocious spouting may indicate the presence of active meristematic cells and low levels of physiological dormancy in the donor material . It may also reflect higher endogenous auxin concentrations, which are known to stimulate early bud break and organogenesis . In addition, the species colonizes gallery forests. In contrast, the significantly delayed sprouting observed in Burkea africana (14 weeks) diverges markedly from previously reported values for other native woody species. For instance, reported much shorter sprouting times of 4, 5, 6 and 8 weeks, respectively, for Bombax costatum, Detarium microcarpum, Vitex doniana, Ximenia americana and Amblygonocarpus andongensis. Several factors may contribute to this divergence: high levels of endogenous abscisic acid (ABA), which suppresses bud growth; structural dormancy related to the thickness or lignification of root tissues; or environmental stress experienced during the collection or handling of cuttings . Beyond species differences, sprouting time is strongly modulated by technical and environmental factors. The timing of cutting collection, the age and position of the donor branch, and the type of substrate used for propagation all play pivotal roles. Additionally, the application of exogenous plant growth regulators is not in the rest.
Comparable results to those of Uvaria chamae have been reported under similar agroecological conditions. identified optimal substrates for Vitex doniana and Ximenia americana as black soil/sawdust mixtures. Similarly, found that the substrate type had no significant effect on Daniellia oliveri, consistent with the findings for Uvaria chamae. In contrast, the results for Burkea africana deviate from previous studies conducted in the Guinean Savannah Highlands of Adamawa, where black soil was identified as the most effective substrate. These findings suggest that the use of porous substrates enhances the circulation of water and air around the cuttings, promoting better budding by maintaining hydration and preventing hypoxic conditions. Substrates offering good aeration and moderate moisture retention typically result in higher budding rates compared to compact or waterlogged media .
These findings support earlier finding by who observed that 20 cm cuttings of Vitex doniana and Detarium microcarpum produced significantly higher bud emergence about five times greater compared to 10 cm cuttings. Likewise, several studies have shown that cuttings between 15 and 20 cm offer the best propagation results. This has been reported respectevely for Faidherbia albida, Spathodea campanulata, Ximenia americana, and Amblygonocarpus andongensis . The superior performance of longer cuttings in tree species has been attributed to their higher reserves of soluble sugars and starches, which provide energy and carbon skeletons essential for cell division and differentiation during bud formation. Longer cuttings, with greater carbohydrate stores, have been shown to root more successfully than shorter ones, as demonstrated in Vitex doniana and Ximenia americana . These carbohydrate reserves also enhance auxin transport and sustain meristematic activity at the cutting base, promoting rooting and bud emergence. postulated that longer cutting segments maintain a higher concentration of stored photosynthates and phytohormones, which facilitate cellular division and differentiation necessary for successful propagation. In contrast, shorter cuttings are more susceptible to desiccation and may lack sufficient energy reserves, thereby limiting their capacity for bud initiation and root development. The biochemical basis underlying these observations is further supported by studies indicating a positive correlation between root elongation and endogenous sugar concentrations .
The non-significant interaction of substrate × cutting length for shoot formation was not jointly influenced by these two factors. This suggests that substrate conditions and cutting length acted independently. Shoot formation is primarily controlled by endogenous reserves and bud viability, while the substrate mainly provides suitable moisture. Similar result was observed by on Amblygonocarpus andongensis.
The relationship between cutting length and shoot production ois consistent with the findinds of for Detarium microcarpum, as well as those of and for Vitex doniana and Securidaca longepedunculata in the Guinean savannah highlands. These authors reported a positive correlation between cutting size and the number of aerial shoots produced. This phenomenon may be attributed to the presence of carbohydrates in the cuttings, which could potentially exert an influence on the growth and development of the aerial shoots.
The results obtained for shoot height suggest that substrate type exerted minimal influence on shoot elongation in the species studied under the experimental conditions. This observation may reflect a conservative growth strategy in B. africana, or alternatively, a reduced sensitivity to substrate composition during early developmental stages. These findings imply that substrate variation did not substantially modulate shoot growth in these taxa. Notably, our results contrast with those reported by , who found a significant effect of substrate type on shoot height. Such divergence may point to considerable interspecific or intraspecific variability, potentially attributable to genetic differences, microenvironmental heterogeneity
The result observed for Burkea africana on the number of leaves indicating that the black soil alone provided more favorable conditions for leaf development in this species. This result may reflect differences in nutrient availability, substrate texture, or moisture retention between the two treatments. The results observed from Uvaria chamae may be due to the large variability in this species can be attributed to intrinsic biological variability or heterogeneous responses to environmental conditions. Overall, these findings suggest species-specific responses to substrate composition, with B. africana showing a clear preference for black soil, while U. chamae did not exhibit statistically significant differences in leaf production across treatments.
4.2.2. Rooting
Following the emergence of young leaves formed via photosynthetic activity, auxins and carbohydrates are translocated through a complex vascular system to the basal portion of the rootstock cuttings (RSC), thereby initiating rhizogenesis. In the present study, the rooting process was found to be significantly slower than the sprouting process, consistent with the findings of , who reported that root initiation generally requires more time than bud emergence. Among the various substrates tested, the black soil and sawdust mixture resulted in the highest rooting rate, corroborating the findings of . The use of mixed substrates appears to offer a more effective medium for root development. Comparable results were reported by for Dalbergia melanoxylon in Tanzania, where a sand and compost mixture outperformed sand alone in promoting root formation. No adventitious roots were observed in Burkea africana, a result that aligns with observations by in Diospyros mespiliformis cuttings in Benin, where no root formation occurred after 29 weeks. This lack of rooting may be attributed to substrate characteristics, particularly porosity, which facilitates root penetration and elongation. Porous substrates have been shown to reduce compaction, thereby enhancing root proliferation . Furthermore, such substrates provide a balance between moisture retention and drainage, ensuring optimal water availability for root initiation and development. The absence of adventitious root formation in Burkea africana root cuttings may be attributed to the physiological age of the donor plants, as the cuttings were obtained from mature trees. Similarly, Bisongin et al. (2020) reported that root system regeneration did not occur in root cuttings derived from adults Cordia trichotoma trees, whereas successful rooting was on served in material collected from juvenile plants.
The results obtain in Burkina Faso on Detarium microcarpum revealed divergent pattern compared to those of the present study. The observed influence of cutting length on rooting performance is consistent with the hypothesis that longer cuttings possess greater reserves of carbohydrates, nutrients, and endogenous auxins, which collectively support adventitious root formation. Longer segments may also maintain higher cambial activity and a more favorable hormonal balance, both of which are critical for root induction . In addition, sucrose has been shown to play a pivotal role in promoting adventitious rooting, not only as an energy source but also by facilitating auxin transport and stimulating cell division . These results are in agreement with findings reported by Binwe et al. (2024), who demonstrated that cutting length significantly affected the rooting success of Ximenia americana root cuttings. This suggests that the positive influence of cutting length may be a common feature across diverse tropical species, although the magnitude of the response may vary depending on species-specific physiological traits and environmental interactions.
The combination of black soil and sawdust produced the greatest root length per plant among the tested substrates. This outcome is likely due to the reduced bulk density of the black soil/ sawdust mixture which enhances soil aeration and reduces mechanical resistance to root growth. Such physical properties facilitate easier root penetration and elongation, thereby supporting the development of longer root systems. This observation aligns with the findings of , who emphasized that effective rooting substrates are those specifically prepared to provide optimal conditions namely, freedom from pests and pathogens, adequate air-filled porosity, sufficient moisture retention, and a suitable bulk density conducive to root development.
4.3. Carbohydrate Content on Developmental Stages of Cuttings
The results of the principal component analysis (PCA) highlight distinct associations between carbohydrate fractions and the developmental stages of cuttings in Uvaria chamae and Burkea africana. In U. chamae, the total variance explained by the first two axes (100%) indicates a strong structuring of the data around carbohydrate content and cutting response. The strong positive correlation of soluble sugars and sucrose with F1, and their spatial proximity to budded cuttings, suggests a significant role of these sugars in bud activation and organogenesis . This finding supports previous studies showing that readily available sugars can act not only as energy sources but also as signaling molecules in the initiation of bud outgrowth . Interestingly, rooted cuttings in U. chamae were located away from all carbohydrate vectors, suggesting a more complex or indirect relationship between carbohydrate content and root formation . This pattern aligns with observations that high carbohydrate levels can inhibit rooting in some species, possibly due to hormonal imbalances or insufficient carbohydrate mobilization toward the root initiation zone. Control cuttings, which showed no visible development, were closely associated with starch. This may indicate a lack of metabolic activation, where carbohydrates remain stored rather than being mobilized for growth. Starch accumulation in inactive tissues could reflect physiological dormancy or low metabolic demand, consistent with their non-responsive status. For B. africana, the one-dimensional PCA structure (100% variance explained by F1) suggests a simpler or more uniform relationship between carbohydrate fractions and cutting development. The location of budded cuttings near soluble sugars on the negative side of F1 further reinforces the idea that soluble carbohydrates support early developmental processes. Conversely, control cuttings were aligned with starch and sucrose on the positive side of the axis, again suggesting that limited mobilization of these carbohydrates may be associated with developmental arrest . In general, the budding and rooting rates were generally negatively correlated with carbohydrate content, with rooting rate showing particularly strong negative associations with starch indicating its use as energy during rooting process.
5. Conclusion
This study provides important evidence that Burkea africana and Uvaria chamae, two native species of the guinean savannah highlands of Adamawa, possess the potential for vegetative propagation through root segment cuttings. The successful induction of budding and shoot development in both species marks a promising step toward their domestication and wider utilization in agroforestry systems.
The success of propagation was shown to be contingent upon a combination of biological, environmental, and technical factors. Among these, seasonal variation in carbohydrate reserves emerged as a critical determinant, with the onset of the rainy season identified as the most favorable period for RSC collection. This phase coincided with higher concentrations of starch, soluble sugars, and sucrose in the root tissues, enhancing the regenerative capacity of the cuttings.
Among the substrates tested, the black soil/sawdust mixture demonstrated superior performance for Uvaria chamae, significantly enhancing both the budding rate and aerial shoot formation. For Burkea africana, black soil alone was found to be the most suitable substrate. Moreover, a cutting length of 20 cm consistently produced the highest results across multiple parameters, including budding rate and root development.
Despite these successes, Burkea africana did not form adventitious roots under the experimental conditions, indicating a potential limitation or the need for further refinement in propagation techniques for this species. Soluble sugars and sucrose are associated with bud activation, while starch accumulation correlates with developmental arrest in both Uvaria chamae and Burkea africana. Root formation in U. chamae appears to involve more complex carbohydrate dynamics. These findings highlight the species-specific responses to vegetative propagation technics and underscore the importance of tailoring protocols to individual plant requirements.
Abbreviations

RSC

Root Segment Cuttings

Acknowledgments
The authors are deeply grateful to the anonymous reviewers for their valuable comments, which significantly improved the quality of the manuscript.
Author Contributions
Jean de Dieu Wangbitching: Conceptualization, Resources, Writing – original draf, Software, Writing – review & editing
Youhana Dangai: Data curation, Methodology,Resources
Haman Zepherin Oumarou: Formal Analysis, Investigation, Methodology
Guidawa Fawa: Methodology,Validation,Ressources
Jean-Baptiste Binwe: Data curation, Methodology, Software
Rodrigue Damba Ameti Madi: Data curation, Methodology, Investigation
Herve Joseph Ewodo Apana: Data curation, Methodology, Investigation
Floriane Sorelle Eyenga: Data curation, Methodology, Investigation
Pierre Marie Mapongmetsem: Validation, Resources, Data Curation
Conflicts of Interest
The authors declare no conflict of interest.
References
[1] Neya, B., Hakkou, M., Pétrissans, M., & Gérardin, P., (2004). On the durability of Burkea africana heartwood: evidence of biocidal and hydrophobic properties responsible for durability. Annals of Forest Science, 61(3): 277-282
[2] Mulofwa, J., Simute, S., & Tengnäs, B., (1994). Agroforestry: manual for extension workers in Southern Province, Zambia.
[3] Ezenyi, I. C., Okpoko, C. K., Ufondu, C. A., Okhale, S. E., & Adzu, B. (2021). Antiplasmodial, antinociceptive and antipyretic potential of the stem bark extract of Burkea africana and identification of its antiplasmodial-active fraction. Journal of traditional and complementary medicine, 11(4): 311-317.
[4] Omulen L., Acere-lervick K., Ndyabarema R., Tumwijukyea, Asio S., (1997). District environment profile: Bushenyi. NORPLAN and Bushenyi District. Internal Report, 100 p.
[5] Meunier Q., Bellefontaine R., Monteuuis O., (2008). Vegetative propagation of medicinal trees and shrubs for the benefit of rural communities in Uganda. Bois et Forêts des Tropiques, 295(2): 71-82.
[6] Moupela, C., Doucet, J. L., Daïnou, K., Meunier, Q., & Vermeulen, C. (2013). Propagation trials by seed and air layering of Coula edulis Baill., and prospects for its domestication. Bois et Forêts des Tropiques., 318(4), 3-13
[7] Bellefontaine R., Meunier Q., Ichaou A., Morin A., Mapongmetsem P.M., Belem B., Azihou F., Houngnon A., Abdourhamane H., 2018. Regeneration by seeds and low-cost vegetative propagation (suckers and root segment cuttings) CIRAD. 460 p.
[8] Ouédraogo E., Mando A., & Brussaard L., 2004. Soil macrofaunal-mediated organic resource disappearance in semi-arid West Africa. Applied Soil Ecology, 27(3): 259-267.
[9] Bellefontaine R., Monteuuis O., (2002). Le drageonnage des arbres hors forêt: un moyen pour revégétaliser partiellement les zones arides et semi-arides sahéliennes ? In Verger M. (Ed) Multiplication végétative des ligneux forestiers, fruitiers et ornementaux. Montpellier, France: Cirad-Inra. Cirad
[10] Hannah J., Jan Beniest., (2003). La multiplication végétative des ligneux en agroforesterie. Manuel de formation et bibliographie,162 p.
[11] Mapongmetsem P.M., Njomba E., Fawa G., Oumarou Z., Dangai Y., Bellefontaine R., (2017). Vegetative Propagation of Vitex doniana Sweet from Root Segments Cuttings: Effects of Substrate and Length of Cuttings on the Rooting Ability. Annals of Experimental Biology, 5(1): 18-24.
[12] Letouzey R., (1968). Phytogéographie du Cameroun. Edition Lechevalier, 518p.
[13] Ministère de l’Environnement et des Forêts (MINEF), (1994). Diagnosis of the environment. Rio World Summit;.p. 113.
[14] Yonkeu S., 1983. Végétation des pâturages de l’Adamaoua (Cameroun): écologie et potentialités pastorales. Thèse de Doctorat. Univ. Rennes I, France. 207p.
[15] Mapongmetsem P.M., Alium P.S., Raouguedam J., Bawa K.L., Fawa G., (2016a). Vegetative propagation of Sclerocarya birrea, (A. Rich.) Hochst. from root segments cuttings: effect of substrate and root diameter. Annals of Experimental Biology, 4(2): 23-32.
[16] Miller, G. L. (1972). Use of dinitrosalicylic acid reagent for determination of reducing sugars. Analytical Chemistry, 31, 426–428.
[17] Dubois, M., Gilles, K. A., Hamilton, J. K., Rebers, P. A., & Smith, F. (1956). Colorimetric method for determination of sugars and related substances. Analytical Chemistry, 28(3), 350–356.
[18] Jarvis, C. E., & Walker, J. R. (1993). Simultaneous, rapid, spectrophotometric determination of total starch, amylose and amylopectin. Journal of the Science of Food and Agriculture, 63(1): 53-57.
[19] Ky-Dembélé C., Tigabu M., Bayala J., Savadogo P., Boussim I.J., Oden P.C., (2010). Clonal propagation of Detarium microcarpum from root cuttings. Silva Fennica, 44(5): 775-787.
[20] Mapongmetsem P. M., Fawa G., Noubissie-Tchiagam J. B., Nkongmeneck B. A., Biaou S. S. H., Bellefontaine R., (2016 b). Vegetative propagation of Vitex doniana Sweet from root segments cuttings. Bois et Forêts des Tropiques, 327(1): 29 – 37.
[21] Oumarou Z.H., Hamaya Y., Tsobou R., Abdoulaye H., Bellefontaine R., Mapongmetsem P.M., (2018). Vegetative propagation of Securidaca longepedunculata Fresen by cutting root segments. Afrique Science, 14(6): 388-399.
[22] Stenvall, N., Piisilä, M., & Pulkkinen, P. (2009). Seasonal fluctuation of root carbohydrates in hybrid aspen clones and its relationship to the sprouting efficiency of root cuttings. Canadian Journal of Forest Research, 39(8), 1531-1537.
[23] Sami, F., Yusuf, M., Faizan, M., Faraz, A., & Hayat, S., 2016. Role of sugars under abiotic stress. Plant physiology and biochemistry, 109, 54-61.
[24] Coates palgrave, K (2002). Tree of southern Africa.Struik, Cape town,1212 pp.
[25] Frost, P. (1996). The ecology of miombo woodlands. The miombo in transition: Woodlands and welfare in Africa, 266, 11-57.
[26] Burkill, H. M. (1985). The Useful Plants of West Tropical Africa. Volume 1, Families A–D (2nd edition). Royal Botanic Gardens, Kew. Vol. 1: 960 p.
[27] Landhäusser, S. M., & Lieffers, V. J., (2003). Seasonal changes in carbohydrate reserves in mature northern Populus tremuloides clones. Trees, 17(6): 471-476.
[28] Oumarou H. Z., Hamawa Yougouda, Tsobou R., Dangai Yohana, Binwe J. B., Madi Amedi Damba R., Abdoulaye Herbert, Wangbitching J. D. D., Fawa Guidawa, Mapongmetsem P. M., (2019). Vegetative Propagation of Bombax costatum Pellegr. & Vuillet (Malvaceae) by Root Segments Cuttings: Effects of Mother Tree Diameter and Origin of Cuttings. American Journal of Agriculture and Forestry, 7(6): 248-258.
[29] Zida, D., Tigabu, M., Sawadogo, L., Tiveau, D., & Odén, P. C. (2009). Long‐term effects of prescribed early fire, grazing and selective tree cutting on seedling populations in the Sudanian savanna of Burkina Faso. African Journal of Ecology, 47(1), 97-108.
[30] Krishnamurthy, Kulithalai & Bahadur, Bir & Adams, Sebastian & Venkatasubramanian, Padma. (2015). Meristems and Their Role in Primary and Secondary Organization of the Plant Body. 151p.
[31] Vanneste, S., Pei, Y., & Friml, J. (2025). Mechanisms of auxin action in plant growth and development. Nature Reviews Molecular Cell Biology, 26(10): 648–666.
[32] Ky-Dembele, Catherine & Traoré, Fatoumata & B., Koné & Bayala, Jules & Antoine, Kalinganire & Bonneville, Jean & Olivier, Alain. (2015). Le bouturage est-il une option envisageable au Sahel ? Sahel Agroforesterie. 20. 4-5.
[33] Belem B., Boussim J.I., Bellefontaine R., Guinko S., (2008). Stimulation du drageonnage de Bombax costatum Pelegr. et Vuillet par blessures de racines au Burkina Faso. Bois et Forêts des Tropiques, 295(1): 71-79.
[34] Binwe J.B., Hamawa Y., Wangbitching J.D.D., Madi A.D.R., Apana E.J.H., Oumarou H.Z., Abdoulaye H., Fawa G., Mapongmetsem P. M., (2024). Influence of substrate and length on the ability of root segments cuttings of Ximenia americana L. to regenerate. International Journal of Research in Agronomy 7(9): 106-113.
[35] Wangbitching J. D D, Yougouda, H., Guidawa, F., Binwe, J. B., Madi Ameti Damba R., Apana Ewodo H. J., Wamba Sopgou D.P., Oumarou H. Z., Abdoulaye H., & Mapongmetsem P. M. P., 2024. Influence of Substrate and Length on The Ability of Root Segments Cuttings of Amblygonocarpus andongensis (Welw. ex Oliv.) Exell & Torre to Regenerate. Discoveries in Agriculture and Food Sciences, 12(6): 56-71.
[36] Rowe, J.H., Topping, J.F., Liu, J. and Lindsey, K., 2016, Abscisic acid regulates root growth under osmotic stress conditions via an interacting hormonal network with cytokinin, ethylene and auxin. New Phytologist, 211: 225-239.
[37] Houehounha R., Avohou H.T., Sinsin B., Tandjiekpon A.M., (2009). Approches de régénération artificielle de Daniellia oliveri (Rolfe) Hutchison et Dalziel. International Journal of Biological and Chemical Sciences, 3(1): 7-19
[38] Hartmann H.T., Kaster D.E., Davies F.T., Geneve R.L., (2004). Plant Propagation: Principles and Practices. 6th ed. Prentice Hall of India Private Limited, New Delhi, India, p.770.
[39] Harivel A., Bellefontaine R., Ousmane B., (2006). Aptitude à la multiplication végétative de huit espèces forestières d‟intérêt au Burkina Faso. Bois et Forêts des Tropiques, 288(2): 39-50.
[40] Leakey, R.R., & Simons, A.J. (1997). The domestication and commercialization of indigenous trees in agroforestry for the alleviation of poverty. Agroforestry Systems, 38, 165-176.
[41] Robinson J.C., Schwabe W.W., 1977. Studies on regeneration of Apple cultivars from root cuttings. Propagation aspects. Journal of Horticultural Science, 52(2): 205–220.
[42] Stenvall N., Haapala T., PulkkinenP., (2004). Effect of genotype, age and treatment of stock plants on propagation of hybrid aspen (Populus tremula x Populus tremuloides) by root cuttings. Scandinavian Journal of Forest Research, 19(4): 303–311.
[43] Magingo, F. S. S., & McP. Dick, J. (2001). Propagation of two miombo woodland trees by leafy stem cuttings obtained from seedlings. Agroforestry systems, 51(1), 49-55.
[44] Fagge A. A. & Manga A. A., (2011). Effect of sowing media and gibberellic acid on the growth and seedling establishment of Bougainvillea glabra, Ixora coccinea and Rosa chinensis. 2. Root Characters. Bayero Journal of Pure and Applied Sciences, 4(2): 155-159.
[45] Bisognin, D. A., Kielse, P., Lencina, K. H., & Mello, U. S. D. (2020). Vegetative propagation of Cordia trichotoma (Vell.) arrab. ex steud. by cuttings from shoots and roots. Cerne, 26, 265-271.
[46] Da Costa, C. T., de Almeida, M. R., Ruedell, C. M., Schwambach, J., Maraschin, F. S., & Fett-Neto, A. G. (2013). When stress and development go hand in hand: Main hormonal controls of adventitious rooting in cuttings. Frontiers in Plant Science, 4, 133.
[47] Thorpe, T. A. 2007. History of plant tissue culture. Molecular Biotechnology, 37(2): 169–180
[48] Eveland, A. L., & Jackson, D. P. (2012). Sugars, signalling, and plant development. Journal of experimental botany, 63(9), 3367-3377.
[49] Wojtusik T., Boyd M. T. and Felker P., 1994: Effects of Different Media on Vegetative Propagation of Prosopic Cuttings Under Solar Power. Journal of Forest Ecology and Management, 69(1-3): 26-71.
[50] Rosa M., Prado C., Podazza G., Interdonato R., González J.A., Hilal M., Prado F.E., (2009). Soluble sugars--metabolism, sensing and abiotic stress: a complex network in the life of plants. Plant Signal Behav; 4(5): 388-93
[51] Ciereszko, I. (2018). Regulatory roles of sugars in plant growth and development. Acta Societatis Botanicorum Poloniae, 87(2).
[52] Bartolini G., Pestelli P., Toponi M. A. & Di monte G., 1996. Rooting and carbohydrate availability in Vitis 140 Ruggeri stem cuttings. Vitis, 35(1): 11–14.
[53] MacNeill, G. J., Mehrpouyan, S., Minow, M. A., Patterson, J. A., Tetlow, I. J., & Emes, M. J. (2017). Starch as a source, starch as a sink: the bifunctional role of starch in carbon allocation. Journal of experimental botany, 68(16): 4433-4453.
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    Wangbitching, J. D. D., Dangai, Y., Oumarou, H. Z., Fawa, G., Binwe, J., et al. (2026). Interactive Effects of Substrate and Length on the Ability of Root Segment Cuttings of Burkea Africana (Hook) and Uvaria Chamae (P. Beauv) to Regenerate. American Journal of Agriculture and Forestry, 14(2), 74-91. https://doi.org/10.11648/j.ajaf.20261402.11

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    ACS Style

    Wangbitching, J. D. D.; Dangai, Y.; Oumarou, H. Z.; Fawa, G.; Binwe, J., et al. Interactive Effects of Substrate and Length on the Ability of Root Segment Cuttings of Burkea Africana (Hook) and Uvaria Chamae (P. Beauv) to Regenerate. Am. J. Agric. For. 2026, 14(2), 74-91. doi: 10.11648/j.ajaf.20261402.11

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    AMA Style

    Wangbitching JDD, Dangai Y, Oumarou HZ, Fawa G, Binwe J, et al. Interactive Effects of Substrate and Length on the Ability of Root Segment Cuttings of Burkea Africana (Hook) and Uvaria Chamae (P. Beauv) to Regenerate. Am J Agric For. 2026;14(2):74-91. doi: 10.11648/j.ajaf.20261402.11

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  • @article{10.11648/j.ajaf.20261402.11,
      author = {Jean de Dieu Wangbitching and Youhana Dangai and Haman Zepherin Oumarou and Guidawa Fawa and Jean-Baptiste Binwe and Rodrigue Damba Ameti Madi and Herve Joseph Ewodo Apana and Floriane Sorelle Eyenga and Pierre Marie Mapongmetsem},
      title = {Interactive Effects of Substrate and Length on the Ability of Root Segment Cuttings of Burkea Africana (Hook) and Uvaria Chamae (P. Beauv) to Regenerate},
      journal = {American Journal of Agriculture and Forestry},
      volume = {14},
      number = {2},
      pages = {74-91},
      doi = {10.11648/j.ajaf.20261402.11},
      url = {https://doi.org/10.11648/j.ajaf.20261402.11},
      eprint = {https://article.sciencepublishinggroup.com/pdf/10.11648.j.ajaf.20261402.11},
      abstract = {The Guinean Savanah Highlands of Adamawa is replete with multipurpose tree species, among which Burkea africana and Uvaria chamae are particularly noteworthy. Despite their importance, they remain in the wild and are subjected to overexploitation. The present study aims to contribute to the domestication of these species by root segment propagation. Specifically, the study aims to evaluate the seasonal variations in carbohydrate reserves (starch, soluble sugars, and sucrose) in other to determine the most favorable period for root cutting collection, assess the effect of substrate and length of root segment cuttings on the budding and rooting capacity of these species, evaluate the effect of carbohydrate content on bud emergence, root formation, callus induction, and control response in cuttings. For the seasonal fluctuation of carbohydrate, the experimental design was a complete randomized design with one factor represented by the season, and two replications. In the case the root propagation, the experimental design was a split-plot with three replications. The main treatment comprised three substrates (sand/sawdust, black soil/sawdust, black soil), while the sub-treatments were represented by three lengths of root segments cuttings (RSC) (10, 15, 20cm). The experimental unit consisted of 10 cuttings. Results showed that the onset of the rainy season coincides with peak of starch, soluble sugars and sucrose for Burkea africana and Uvaria chamae, marking the most favorable period for root cutting collection. The budding rate of Burkea africana showed significant variation (0.04 Uvaria chamae the best substrate was the mixture of black soil/sawdust (63.33 ± 45.27%). The number of leaves for Burkea africana was substantially higher in black soil (4.27 ± 2.16). The difference was statistically significant (0.008 Burkea africana and Uvaria chamae, the optimal cutting length for budding was 20 cm (11.11 ± 10.52%, 81.11 ± 26.20%). The rooting rate of Uvaria chamae exhibited considerable variability the best rate was those of the black soil/sawdust substrate (22.22 ± 20.33%). The rooting rate varied from 5.55 ± 4.26% for cuttings of 10 cm to 23.33 ± 21.79% for those of 20 cm.Budded cuttings clustered with soluble sugars and sucrose. Rooted cuttings correlate negatively with all carbohydrate. Control cuttings were closely associated with starch. All these informations are important to develop scales and strategies toward the domestication of this species.},
     year = {2026}
    }
    

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  • TY  - JOUR
    T1  - Interactive Effects of Substrate and Length on the Ability of Root Segment Cuttings of Burkea Africana (Hook) and Uvaria Chamae (P. Beauv) to Regenerate
    AU  - Jean de Dieu Wangbitching
    AU  - Youhana Dangai
    AU  - Haman Zepherin Oumarou
    AU  - Guidawa Fawa
    AU  - Jean-Baptiste Binwe
    AU  - Rodrigue Damba Ameti Madi
    AU  - Herve Joseph Ewodo Apana
    AU  - Floriane Sorelle Eyenga
    AU  - Pierre Marie Mapongmetsem
    Y1  - 2026/03/10
    PY  - 2026
    N1  - https://doi.org/10.11648/j.ajaf.20261402.11
    DO  - 10.11648/j.ajaf.20261402.11
    T2  - American Journal of Agriculture and Forestry
    JF  - American Journal of Agriculture and Forestry
    JO  - American Journal of Agriculture and Forestry
    SP  - 74
    EP  - 91
    PB  - Science Publishing Group
    SN  - 2330-8591
    UR  - https://doi.org/10.11648/j.ajaf.20261402.11
    AB  - The Guinean Savanah Highlands of Adamawa is replete with multipurpose tree species, among which Burkea africana and Uvaria chamae are particularly noteworthy. Despite their importance, they remain in the wild and are subjected to overexploitation. The present study aims to contribute to the domestication of these species by root segment propagation. Specifically, the study aims to evaluate the seasonal variations in carbohydrate reserves (starch, soluble sugars, and sucrose) in other to determine the most favorable period for root cutting collection, assess the effect of substrate and length of root segment cuttings on the budding and rooting capacity of these species, evaluate the effect of carbohydrate content on bud emergence, root formation, callus induction, and control response in cuttings. For the seasonal fluctuation of carbohydrate, the experimental design was a complete randomized design with one factor represented by the season, and two replications. In the case the root propagation, the experimental design was a split-plot with three replications. The main treatment comprised three substrates (sand/sawdust, black soil/sawdust, black soil), while the sub-treatments were represented by three lengths of root segments cuttings (RSC) (10, 15, 20cm). The experimental unit consisted of 10 cuttings. Results showed that the onset of the rainy season coincides with peak of starch, soluble sugars and sucrose for Burkea africana and Uvaria chamae, marking the most favorable period for root cutting collection. The budding rate of Burkea africana showed significant variation (0.04 Uvaria chamae the best substrate was the mixture of black soil/sawdust (63.33 ± 45.27%). The number of leaves for Burkea africana was substantially higher in black soil (4.27 ± 2.16). The difference was statistically significant (0.008 Burkea africana and Uvaria chamae, the optimal cutting length for budding was 20 cm (11.11 ± 10.52%, 81.11 ± 26.20%). The rooting rate of Uvaria chamae exhibited considerable variability the best rate was those of the black soil/sawdust substrate (22.22 ± 20.33%). The rooting rate varied from 5.55 ± 4.26% for cuttings of 10 cm to 23.33 ± 21.79% for those of 20 cm.Budded cuttings clustered with soluble sugars and sucrose. Rooted cuttings correlate negatively with all carbohydrate. Control cuttings were closely associated with starch. All these informations are important to develop scales and strategies toward the domestication of this species.
    VL  - 14
    IS  - 2
    ER  - 

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    1. 1. Introduction
    2. 2. Materials and Methods
    3. 3. Results
    4. 4. Discussion
    5. 5. Conclusion
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