# Impact of environment and genotype-by-environment interaction on functional properties of amylose-free and wild-type cassava starches

Amanda Karlström, John Eiver Belalcazar, Teresa Sánchez, Jorge Ivan Lenis, John Larry Moreno, Monica Pizarro, Julien Ricci, Dominique Dufour, Thierry Tran, Hernan Ceballos

► To cite this version:

Amanda Karlström, John Eiver Belalcazar, Teresa Sánchez, Jorge Ivan Lenis, John Larry Moreno, et al. Impact of environment and genotype-by-environment interaction on functional properties of amylose-free and wild-type cassava starches. Starch/Stärke, 2019, 71 (1-2), pp.1700278. 10.1002/star.201700278. hal-05173590

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# Impact of Environment and Genotype-by-Environment Interaction on Functional Properties of Amylose-Free and Wildtype Cassava Starches

Amanda Karlström, John Belalcazar, Teresa Sánchez, Jorge I. Lenis, John L. Moreno, Mónica Pizarro, Julien Ricci, Dominique Dufour, Thierry Tran, and Hernán Ceballos*

Ten and thirty cassava genotypes producing wildtype and amylose-free (waxy) starch respectively, are grown at low (LA) and intermediate (IA) altitudes (<213 and >900 m above sea level (m.a.s.l.) respectively). The functional properties of the starch from these materials are analyzed. The relative importance of environment, clones (within starch type) and genotype-by-environment interaction (GxE) is assessed through an analysis of variance. GxE effects are non-significant, whereas environment and clones (within starch type) are highly significant in most of the cases (particularly for waxy starches). Results confirm that waxy cassava starches consistently show higher peak viscosity (PV) compared with wildtypes. Peak viscosity is statistically higher in plants grown in IA compared with those from the same genotypes grown in LA, both for waxy and wildtype genotypes. In a second experiment, a single waxy genotype is grown in five locations ranging from 934 to 1485 m.a.s.l. PV is highest in the location with higher altitude. The second highest PV is measured in starches from the second highest location at 1071 m. Both experiments clearly indicate that PV is positively correlated with altitude above sea level where cassava is grown. Altitude above sea level influenced starch granule size: the higher the altitude the larger the average granule size. Waxy starch granules tend to be larger than those from wildtype genotypes.

A. Karlström, J. Belalcazar, T. Sánchez, J. I. Lenis, J. L. Moreno, M. Pizarro, Dr. D. Dufour, Dr. T. Tran, Dr. H. Ceballos
Centro Internacional de Agricultura Tropical (CIAT)
CGIAR Research Program on Roots Tubers and Bananas (RTB)
Programa de Yuca (CIAT), Apartado Aéreo 6713, Cali, Palmira, Colombia
E-mail: h.ceballos@cgiar.org

A. Karlström
Swedish University of Agricultural Sciences
Alnarp, Sweden

J. Ricci, D. Dufour, T. Tran
Centre de Coopération Internationale en Recherche Agronomique pour le Développement (CIRAD)
UMR Qualisud
Montpellier, France

© 2018 The Authors. *Starch* - *Stärke* Published by Wiley Periodicals, Inc.. This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

DOI: 10.1002/star.201700278

## 1. Introduction

Cassava is the second most important source of starch worldwide.[1] About 74–85% of the dry root weight of cassava is starch. Compared with other tropical root and tuber crops, cassava starch and its biosynthesis have been well studied and described.[2] The starch granules are generally round (oval), with a flat surface on one side (truncated). The size of individual granules ranges from 5 to about 40 µm, with reported averages varying from 5.4 to 17.2 mm. A mutation with considerably smaller starch granule size has been reported.[3] Average amylose content in more than 4000 genotypes was 20.7%[4] and several amylose-free mutations have been reported.[5-7] Cassava roots have little fat and proteins facilitating the extraction of a starch that has excellent quality, non-cereal taste, and, if properly extracted, a pure white color.[8]

Environmental conditions have large impact on the starch production and starch quality in different crops. Genotype, environment and genotype-by-environment interactions greatly influence different agronomic traits (root and starch yield, carotenoids content, cyanogenic potential, etc.) in cassava as well.[9-12] Similar results were summarized for other crops.[13] The functional properties of starch is intricately linked to its chemistry and the structure and morphology of the granule. The different enzymes involved in starch biosynthesis pathway have been thoroughly studied and described. Environmental stresses influence the expression/activity of these enzymes. Variations in soil moisture and composition (e.g., N-content and salinity), ambient temperature (both high and low), and atmospheric composition have been shown to affect starch functionality and variations in amylose/amylopectin ratio in cereals.[13,14] These studies, however, focused on the impact of environmental effects on functional properties and did not report on the effect of genotype-by-environment interactions. Environmental effects have been reported for starch quality traits in different cereals and grain crops[15-18] and in potato.[19-21] The relative importance of genotype-by-environment interactions on the functional properties of starch in many crops varies widely.[13-22]
<table><caption>**Table 1.** List of 10 wildtype and 30 waxy genotypes from the first experiment.</caption><thead><tr><th colspan="2">Wildtype starch</th><th colspan="6">Amylose-free (waxy) starch</th></tr><tr><th>Code</th><th>Pedigree</th><th>Code</th><th>Pedigree</th><th>Code</th><th>Pedigree</th><th>Code</th><th>Pedigree</th></tr></thead><tbody><tr><td>N1</td><td>GM214-62</td><td>W12</td><td>GM3904-14</td><td>W10</td><td>GM3938-136</td><td></td><td>GM3941-24</td></tr><tr><td>N2</td><td>GM579-13</td><td>W8</td><td>GM3904-54</td><td>W16</td><td>GM3938-187</td><td>W1</td><td>GM4781-3</td></tr><tr><td>N3</td><td>SM2629-36</td><td></td><td>GM3916-19</td><td>W9</td><td>GM3938-229</td><td>W17</td><td>GM4817-1</td></tr><tr><td>N4</td><td>SM2828-28</td><td></td><td>GM3916-21</td><td>W5</td><td>GM3938-88</td><td>W14</td><td>GM4883-1</td></tr><tr><td>N5</td><td>SM2834-31</td><td></td><td>GM3917-114</td><td>W11</td><td>GM3939-143</td><td>W6</td><td>GM4883-3</td></tr><tr><td>N6</td><td>SM3110-15</td><td></td><td>GM3917-124</td><td>W7</td><td>GM3939-165</td><td></td><td>GM5389-20</td></tr><tr><td>N7</td><td>SM3134-5</td><td>W15</td><td>GM3917-63</td><td></td><td>GM3939-229</td><td></td><td>GM5389-46</td></tr><tr><td>N8</td><td>SM3137-40</td><td>W18</td><td>GM3937-106</td><td>W3</td><td>GM3939-272</td><td></td><td>GM5390-22</td></tr><tr><td>N9</td><td>SM3140-2</td><td></td><td>GM3937-111</td><td></td><td>GM3939-289</td><td>W2</td><td>GM5391-2</td></tr><tr><td>N10</td><td>SM3150-17</td><td></td><td>GM3937-63</td><td>W4</td><td>GM3939-37</td><td>W13</td><td>SM3794-1</td></tr></tbody></table>

Data from only 18 waxy genotypes is presented in this article (identified with a "W" code). The first six digits of the pedigree identify the family, and the numbers after the dash indicate different siblings within a family.

In cassava, significant genotype, environment and interaction effects for swelling power, peak viscosity, and cyanogenic potential when growing cassava under normal and water stress conditions have been reported.[9] In Africa, significant genotype by environment interactions were identified for a range of traits: flour and starch solubility, swelling power, water binding capacity, as well as swelling capacity and pH in gari.[23]

As mentioned above, several sources of amylose-free (waxy) cassava starch have been identified and reported.[5-7] The functional properties of waxy cassava starch have been thoroughly characterized.[24,25] Waxy cassava starch from genetically modified sources have been obtained and their functional properties also reported.[26,27] Commercial cassava varieties producing amylose-free starch have been developed in Thailand and Colombia. The impact of the waxy mutation on dry matter content in the roots has been measured and reported.[28]

The present study was conducted to analyze wildtype and amylose-free starch functional properties when cassava clones were grown at different altitudes above sea level and was part of a M.Sc dissertation.[29] Genetic, environmental, and interaction effects were measured.

## 2. Experimental Section

Two experiments were conducted.

### 2.1. First Experiment

#### 2.1.1. Environments

Environments (all in Colombia) with two contrasting altitudes (measured in meters above sea level, m.a.s.l.) were used. Sincelejo (213 m.a.s.l. Sucre Department) and Barranquilla (31 m.a.s.l., Atlántico Department) are at low altitude (LA) conditions, whereas CIAT-Palmira (965 m.a.s.l., Valle del Cauca Department) has intermediate altitude (IA). The latter is where CIAT's headquarters and the root quality laboratory are located.

#### 2.1.2. Germplasm and Root Production

A group of 10 wildtype starch clones were grown simultaneously in Sincelejo and CIAT (Table 1). Also 30 clones producing amylose-free starch were grown in Barranquilla and CIAT (Table 1). Clones grown at low altitude environments were harvested and commercial-size roots shipped by plane to Palmira where starch was extracted, along with those from the clones

<table><caption>**Table 2.** Quality traits in gels derived from wildtype genotypes grown in low (LA) and intermediate (IA) environments.</caption><thead><tr><th rowspan="2">Altitude</th><th colspan="2">Clarity<br>[PC; %]</th><th colspan="2">Swelling<br>[SW; g g<sup>-1</sup>]</th><th colspan="2">Solubility<br>[SO; % db]</th></tr><tr><th>LA</th><th>IA</th><th>LA</th><th>IA</th><th>LA</th><th>IA</th></tr></thead><tbody><tr><td>N1</td><td>54.9</td><td>61.1</td><td>26.3</td><td>30.6</td><td>16.6</td><td>13.3</td></tr><tr><td>N2</td><td>61.8</td><td>57.0</td><td>28.2</td><td>26.4</td><td>11.8</td><td>17.1</td></tr><tr><td>N3</td><td>60.7</td><td>63.7</td><td>29.3</td><td>23.5</td><td>13.9</td><td>13.8</td></tr><tr><td>N4</td><td>54.9</td><td>63.1</td><td>27.1</td><td>25.5</td><td>14.1</td><td>16.2</td></tr><tr><td>N5</td><td>63.9</td><td>62.5</td><td>35.1</td><td>26.3</td><td>12.6</td><td>13.2</td></tr><tr><td>N6</td><td>70.7</td><td>69.0</td><td>36.1</td><td>35.1</td><td>13.8</td><td>12.1</td></tr><tr><td>N7</td><td>68.0</td><td>69.6</td><td>29.9</td><td>32.6</td><td>14.1</td><td>13.0</td></tr><tr><td>N8</td><td>52.5</td><td>66.4</td><td>24.7</td><td>33.6</td><td>14.2</td><td>14.7</td></tr><tr><td>N9</td><td>61.3</td><td>57.6</td><td>26.2</td><td>26.5</td><td>14.1</td><td>14.4</td></tr><tr><td>N10</td><td>70.6</td><td>57.3</td><td>33.8</td><td>27.3</td><td>14.8</td><td>13.1</td></tr><tr><td>Mean</td><td>61.9</td><td>62.7</td><td>29.7</td><td>28.7</td><td>14.0</td><td>14.1</td></tr><tr><td>S Dev</td><td>0.38</td><td>0.38</td><td>0.94</td><td>0.73</td><td>0.36</td><td>0.22</td></tr></tbody></table>

Statistical significance of the sources of variation (probability > F) from ANOVA

<table><tr><td>Env</td><td>0.5799</td><td>0.3847</td><td>0.8631</td></tr><tr><td>Clon</td><td>0.0051</td><td>0.0119</td><td>0.3615</td></tr><tr><td>GxE</td><td>0.9865</td><td>0.7393</td><td>0.7509</td></tr></table>

At the bottom the statistical significances for Environment (Env), Genetic (Clon) and Genotype-by-Environment interactions (GxE) effects are presented.
grown locally. Amylose content was confirmed by the iodine solution test.[5-7]

## 2.2. Second Experiment

### 2.2.1. Environments

Five different environments were used. Four locations were in Valle del Cauca Department of Colombia: Sonso (934 m.a.s.l.); Corpoica-Palmira (1001 m.a.s.l.); CIAT-Palmira (965 m.a.s.l.); and Darien (1485 m.a.s.l.). The remaining environment (Santander de Quilichao) was in Cauca Department and is about 1071 m.a.s.l. These environments were at IA, except for Darien which is considered to be a high altitude (HA) for cassava standards.

### 2.2.2. Germplasm and Root Production

Only one genotypes (AM206-5) was used for the second experiment. This genotype was the first source of amylose-free starch reported in cassava.[5]

### 2.2.3. Starch Extraction

The freshly cut pieces were suspended in tap water and crushed in a 4 L capacity Waring Commercial blender (New Hartford, CT, USA). The slurry was filtered through a 100 µm sieve. The starch

was allowed to settle for 5-6 h. The supernatant decanted off, then the starch was dried in an oven with fan-forced ventilation at 40 °C for 2 days (Thelco Oven Model 28, Precision Scientific Subsidiary of GCA Corporation, Chicago, USA). This methodology has been used with satisfactory results by the root quality laboratory at CIAT for many years. For example, average dry matter content in the wildtype starches ranged from 92.3% to 93.9%, with and average standard deviation (of the two aliquots) of 0.375, resulting in a coefficient of variation of 0.402.

## 2.3. Functional Properties Determinations

### 2.3.1. Paste Clarity (PC)

A 1% db aqueous dispersion of starch was boiled at 97 °C (1000 m.a.s.l.) and shaken thoroughly every 5 min for 30 min. Transmittance was measured after cooling to room temperature at 650 nm. Three repetitions per genotype were used.

### 2.3.2. Swelling Power and Solubility

The swelling power and solubility of the starch samples were determined using a 1.2% db starch paste (0.336 g db starch in 27 664 g distilled water).[29] This was a departure of the 1.5 % db pastes used in previous publications,[4,30] and was made because of lack of separation of the gel phase using the standard procedure. The starch pastes were prepared in the RNA starting at 35 °C for 1 min, increasing to 75 °C at a rate of 6 °C per minute and holding at 75 °C for 2.5 min. During the first minute, the

<table><caption>Table 3. Starch functional properties (from rapid viscoanalyzer) of wildtype genotypes grown in low (LA) and intermediate (IA) environments.</caption><thead><tr><th rowspan="2">Altit.</th><th colspan="2">PT [°C]</th><th colspan="2">PV [cP]</th><th colspan="2">HP [cP]</th><th colspan="2">CP [cP]</th><th colspan="2">BD [cP]</th><th colspan="2">SB [cP]</th><th colspan="2">CS [cP]</th></tr><tr><th>LA</th><th>IA</th><th>LA</th><th>IA</th><th>LA</th><th>IA</th><th>LA</th><th>IA</th><th>LA</th><th>IA</th><th>LA</th><th>IA</th><th>LA</th><th>IA</th></tr></thead><tbody><tr><td>N1</td><td>73.7</td><td>65.7</td><td>500.1</td><td>935.8</td><td>302.1</td><td>490.8</td><td>363.6</td><td>749.3</td><td>198.0</td><td>445.0</td><td>-136.5</td><td>-186.5</td><td>61.5</td><td>258.5</td></tr><tr><td>N2</td><td>71.6</td><td>69.5</td><td>680.8</td><td>717.4</td><td>491.3</td><td>375.9</td><td>641.8</td><td>436.4</td><td>189.5</td><td>341.5</td><td>-39.0</td><td>-281.0</td><td>150.5</td><td>60.5</td></tr><tr><td>N3</td><td>72.6</td><td>67.7</td><td>630.4</td><td>989.6</td><td>452.9</td><td>589.1</td><td>620.4</td><td>875.6</td><td>177.5</td><td>400.5</td><td>-10.0</td><td>-114.0</td><td>167.5</td><td>286.5</td></tr><tr><td>N4</td><td>71.0</td><td>71.8</td><td>637.6</td><td>680.0</td><td>438.6</td><td>382.5</td><td>602.1</td><td>550.5</td><td>199.0</td><td>297.5</td><td>-35.5</td><td>-129.5</td><td>163.5</td><td>168.0</td></tr><tr><td>N5</td><td>72.1</td><td>67.1</td><td>633.6</td><td>905.0</td><td>435.6</td><td>479.0</td><td>590.1</td><td>738.5</td><td>198.0</td><td>426.0</td><td>-43.5</td><td>-166.5</td><td>154.5</td><td>259.5</td></tr><tr><td>N6</td><td>71.8</td><td>67.0</td><td>653.5</td><td>882.1</td><td>409.5</td><td>426.1</td><td>587.5</td><td>669.6</td><td>244.0</td><td>456.0</td><td>-66.0</td><td>-212.5</td><td>178.0</td><td>243.5</td></tr><tr><td>N7</td><td>70.4</td><td>67.2</td><td>705.0</td><td>857.7</td><td>463.5</td><td>427.7</td><td>656.0</td><td>649.2</td><td>241.5</td><td>430.0</td><td>-49.0</td><td>-208.5</td><td>192.5</td><td>221.5</td></tr><tr><td>N8</td><td>70.7</td><td>68.7</td><td>671.4</td><td>831.6</td><td>453.4</td><td>465.6</td><td>623.9</td><td>675.6</td><td>218.0</td><td>366.0</td><td>-47.5</td><td>-156.0</td><td>170.5</td><td>210.0</td></tr><tr><td>N9</td><td>70.4</td><td>68.4</td><td>745.4</td><td>847.2</td><td>531.4</td><td>492.2</td><td>710.4</td><td>711.2</td><td>214.0</td><td>355.0</td><td>-35.0</td><td>-136.0</td><td>179.0</td><td>219.0</td></tr><tr><td>N10</td><td>70.4</td><td>66.6</td><td>677.9</td><td>938.1</td><td>421.9</td><td>485.1</td><td>587.4</td><td>750.6</td><td>256.0</td><td>453.0</td><td>-90.5</td><td>-187.5</td><td>165.5</td><td>265.5</td></tr><tr><td>Mean</td><td>71.5</td><td>67.9</td><td>653.6</td><td>858.4</td><td>440.0</td><td>461.4</td><td>598.3</td><td>680.6</td><td>213.6</td><td>397.1</td><td>-55.3</td><td>-177.8</td><td>158.3</td><td>219.3</td></tr><tr><td>S Dev</td><td>0.21</td><td>0.22</td><td>5.09</td><td>5.78</td><td>5.14</td><td>6.21</td><td>5.38</td><td>7.36</td><td>2.35</td><td>7.55</td><td>3.75</td><td>8.20</td><td>3.4</td><td>8.15</td></tr></tbody></table><table><caption>Statistical significance of the sources of variation (probability > F) from ANOVA</caption><tbody><tr><td>Env</td><td>&lt;.0001</td><td>&lt;.0001</td><td>0.2157</td><td>0.0118</td><td>&lt;.0001</td><td>&lt;.0001</td><td>0.0002</td></tr><tr><td>Clon</td><td>0.2273</td><td>0.1344</td><td>0.0451</td><td>0.0957</td><td>&lt;.0001</td><td>0.0005</td><td>0.0297</td></tr><tr><td>E°C</td><td>0.6467</td><td>0.9785</td><td>0.9607</td><td>0.9592</td><td>0.7442</td><td>0.5294</td><td>0.9560</td></tr></tbody></table>

PT, pasting temperature; PV, peak viscosity; HP, hot paste (through) viscosity; CP, cool paste (end) viscosity; BD, breakdown; SB, setback; CS, consistency. At the bottom the statistical significances for Environment (Env), Genetic (Clon) and Genotype-by-Environment interactions (GxE) effects are presented.
samples were stirred at 960 rpm and thereafter maintained at 160 rpm during the rest of the analysis. The paste was then immediately transferred to 40 mL Falcon tubes, left to cool to room temperature and centrifuged at 7000 rpm for 10 min. The supernatant and sediment was thereafter collected and weighed ($W_{su}$ and $W_{se}$, respectively) and dried in an oven at 105 °C for 24 and 48 h, respectively, after which they were weighed a second time ($D_{su}$ and $D_{se}$). Two replicates of every genotype were taken. The obtained values were thereafter used to calculate three parameters: the concentration of soluble material in the supernatant (solubility), the swelling power and the volume fraction of the dispersed starch phase ($\phi$).

### 2.3.3. Pasting Properties

Hot starch dispersion viscosity profiles were obtained with a Rapid Visco Analyzer model RVA-4 Series (Newport Scientific, Australia). Further details of the methodology are available.[4,30] Four parameters were measured directly: pasting temperature (PT), peak viscosity (PV), hot paste viscosity at the end of the plateau at 90 °C (HP); cool paste viscosity at the end of the cooling step, as temperature reaches 50 °C (CP). With these values, three additional parameters were calculated: breakdown (BD), estimated as PV-HP; setback (SB), estimated as CP-PV; and consistency (CS), estimated as CP-HP.

### 2.3.4. Granule Size and Distribution

Starch granule size and distribution was determined with a laser diffraction particle size analyzer (Malvern Mastersizer 3000, Malvern, UK), equipped with a AERO S measurement cell. Starch samples were suspended in a flow of air prior to their injection into the instrument. Experimental conditions were adjusted for measurements at an obscuration of 3%. Refractive index of $1.600 \pm 0.101$ was set for the starch. Measurements were run in triplicate on each starch samples per genotype at room temperature. Wildtype samples from the low altitude environment were not available for the analysis of granule size. Samples from AM206-5 described in the second experiment were not available either.

### 2.3.5. Statistical Analysis

Statistical analysis was made using the SAS Software and the Proc GLM procedure.[31]

## 3. Results and Discussion

### 3.1. First Experiment

Data for all variables and starch samples was available, except for solubility and swelling power in one of the waxy genotypes. Although 30 waxy starch clones were evaluated (except for granule size) individual data is presented from 18 of them. These 18 genotypes were selected because they properly illustrate the

kind of phenotypic variation (e.g., maximum and minimum values) observed in the entire set of clones and characteristics evaluated. All 30 clones, however, were used in statistical analyses of variance. The 30 waxy starch genotypes were derived from 16 different families (identified by the first six digits in the pedigree). The 18 waxy clones from which data is presented represent 10 of these families. Table 2 and 3 present the results from the ten clones producing wildtype (containing amylose) starch. Equivalent results for the 18 of the 30 waxy starch genotypes are presented in Table 4 and 5.

The altitude above sea level strongly influenced starch granule size. On average the 18 waxy starch samples had a median diameter $Dx(50)$ of $14.41 (\pm 0.78)$ and $15.54 \mu m$ ($\pm 0.89$) for samples from the LA and IA environments, respectively. Paired t-test was statistically significant ($p = 2.49 \times 10^{-10}$). $Dx(50)$ from samples in the LA environment ranged from 13.02 to $15.96 \mu m$, whereas those from IA varied from 14.01 to $17.68 \mu m$. The comparison of wildtype versus waxy starches could be done only

<table><caption>**Table 4.** Quality traits in gels derived from waxy genotypes grown in low (LA) and intermediate (IA) environments.</caption><thead><tr><th rowspan="2">Parameter</th><th colspan="2">Clarity [%]</th><th colspan="2">Swelling [g g<sup>-1</sup>]</th><th colspan="2">Solubility [% db]</th></tr><tr><th>LA</th><th>IA</th><th>LA</th><th>IA</th><th>LA</th><th>IA</th></tr></thead><tbody><tr><td>Altitude</td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>W1</td><td>75.6</td><td>67.7</td><td>n.a.</td><td>n.a.</td><td>n.a.</td><td>n.a.</td></tr><tr><td>W2</td><td>68.6</td><td>67.6</td><td>69.4</td><td>58.1</td><td>6.1</td><td>8.7</td></tr><tr><td>W3</td><td>69.4</td><td>69.4</td><td>63.5</td><td>59.0</td><td>8.6</td><td>5.2</td></tr><tr><td>W4</td><td>74.2</td><td>72.2</td><td>73.8</td><td>69.0</td><td>7.4</td><td>6.2</td></tr><tr><td>W5</td><td>61.2</td><td>65.9</td><td>54.6</td><td>50.5</td><td>8.6</td><td>4.8</td></tr><tr><td>W6</td><td>68.1</td><td>66.7</td><td>55.9</td><td>61.1</td><td>9.8</td><td>5.8</td></tr><tr><td>W7</td><td>67.2</td><td>67.9</td><td>56.4</td><td>53.1</td><td>9.6</td><td>9.5</td></tr><tr><td>W8</td><td>63.7</td><td>55.6</td><td>52.2</td><td>54.2</td><td>7.6</td><td>7.9</td></tr><tr><td>W9</td><td>68.2</td><td>69.8</td><td>62.8</td><td>60.4</td><td>10.5</td><td>9.3</td></tr><tr><td>W10</td><td>65.8</td><td>67.9</td><td>56.9</td><td>60.1</td><td>9.4</td><td>5.7</td></tr><tr><td>W11</td><td>67.9</td><td>70.2</td><td>53.5</td><td>75.4</td><td>7.5</td><td>8.9</td></tr><tr><td>W12</td><td>64.8</td><td>59.1</td><td>55.8</td><td>62.1</td><td>8.4</td><td>6.1</td></tr><tr><td>W13</td><td>67.4</td><td>62.8</td><td>53.6</td><td>70.6</td><td>7.1</td><td>6.8</td></tr><tr><td>W14</td><td>70.5</td><td>66.1</td><td>59.5</td><td>54.0</td><td>8.6</td><td>8.8</td></tr><tr><td>W15</td><td>64.7</td><td>65.6</td><td>59.8</td><td>57.8</td><td>6.7</td><td>5.8</td></tr><tr><td>W16</td><td>69.5</td><td>68.9</td><td>67.4</td><td>62.8</td><td>17.6</td><td>9.4</td></tr><tr><td>W17</td><td>65.7</td><td>71.5</td><td>61.5</td><td>68.7</td><td>14.6</td><td>11.1</td></tr><tr><td>W18</td><td>61.1</td><td>61.7</td><td>59.7</td><td>47.6</td><td>5.2</td><td>5.0</td></tr><tr><td>Min</td><td>60.2</td><td>55.6</td><td>52.2</td><td>47.6</td><td>5.2</td><td>4.1</td></tr><tr><td>Max</td><td>75.6</td><td>72.2</td><td>73.8</td><td>75.4</td><td>17.6</td><td>11.1</td></tr><tr><td>Mean</td><td>66.7</td><td>66.5</td><td>58.8</td><td>61.0</td><td>8.4</td><td>7.3</td></tr><tr><td>S Dev</td><td>0.98</td><td>0.56</td><td>2.84</td><td>3.26</td><td>0.87</td><td>0.92</td></tr></tbody></table>

Statistical significance of the sources of variation (probability > F) from ANOVA

<table><tr><td>Env</td><td>0.5391</td><td>0.1274</td><td>0.0062</td></tr><tr><td>Clon</td><td>&lt;.0001</td><td>0.1698</td><td>&lt;.0001</td></tr><tr><td>GxE</td><td>0.8475</td><td>0.7043</td><td>0.8635</td></tr></table>

At the bottom the statistical significances for Environment (Env), Genetic (Clon) and Genotype-by-Environment interactions (GxE) effects are presented.
<table><caption>**Table 5.** Starch functional properties (from rapid viscoanalyzer) of waxy genotypes grown in low (LA) and intermediate (IA) environments.</caption><thead><tr><th rowspan="2">Altit.</th><th colspan="2">PT [°C]</th><th colspan="2">PV [cP]</th><th colspan="2">HP [cP]</th><th colspan="2">CP [cP]</th><th colspan="2">BD [cP]</th><th colspan="2">SB [cP]</th><th colspan="2">CS [cP]</th></tr><tr><th>LA</th><th>IA</th><th>LA</th><th>IA</th><th>LA</th><th>IA</th><th>LA</th><th>IA</th><th>LA</th><th>IA</th><th>LA</th><th>IA</th><th>LA</th><th>IA</th></tr></thead><tbody><tr><td>W1</td><td>71.0</td><td>66.2</td><td>885.8</td><td>1079.2</td><td>233.8</td><td>336.2</td><td>273.3</td><td>368.2</td><td>652.0</td><td>743.0</td><td>-612.5</td><td>-711.0</td><td>39.5</td><td>32.0</td></tr><tr><td>W2</td><td>70.7</td><td>66.5</td><td>886.0</td><td>1090.7</td><td>257.5</td><td>371.7</td><td>287.0</td><td>395.2</td><td>628.5</td><td>719.0</td><td>-599.0</td><td>-695.5</td><td>29.5</td><td>23.5</td></tr><tr><td>W3</td><td>72.6</td><td>67.6</td><td>1074.1</td><td>1223.4</td><td>447.1</td><td>515.4</td><td>511.1</td><td>596.4</td><td>627.0</td><td>708.0</td><td>-563.0</td><td>-627.0</td><td>64.0</td><td>81.0</td></tr><tr><td>W4</td><td>72.3</td><td>65.7</td><td>1040.4</td><td>1168.1</td><td>494.9</td><td>457.6</td><td>535.4</td><td>496.6</td><td>545.5</td><td>710.5</td><td>-505.0</td><td>-671.5</td><td>40.5</td><td>39.0</td></tr><tr><td>W5</td><td>72.8</td><td>66.1</td><td>1006.8</td><td>1097.7</td><td>458.3</td><td>475.2</td><td>542.3</td><td>552.2</td><td>548.5</td><td>622.5</td><td>-464.5</td><td>-545.5</td><td>84.0</td><td>77.0</td></tr><tr><td>W6</td><td>72.7</td><td>66.8</td><td>962.6</td><td>1148.1</td><td>443.6</td><td>481.1</td><td>504.6</td><td>577.1</td><td>519.0</td><td>667.0</td><td>-458.0</td><td>-571.0</td><td>61.0</td><td>96.0</td></tr><tr><td>W7</td><td>72.0</td><td>66.9</td><td>1071.3</td><td>1080.4</td><td>456.8</td><td>424.4</td><td>520.3</td><td>452.9</td><td>614.5</td><td>656.0</td><td>-551.0</td><td>-627.5</td><td>63.5</td><td>28.5</td></tr><tr><td>W8</td><td>72.0</td><td>67.1</td><td>925.7</td><td>1063.4</td><td>441.7</td><td>444.9</td><td>507.2</td><td>512.4</td><td>484.0</td><td>618.5</td><td>-418.5</td><td>-551.0</td><td>65.5</td><td>67.5</td></tr><tr><td>W9</td><td>71.4</td><td>65.6</td><td>1071.0</td><td>1203.9</td><td>480.0</td><td>478.4</td><td>527.0</td><td>526.9</td><td>591.0</td><td>725.5</td><td>-544.0</td><td>-677.0</td><td>47.0</td><td>48.5</td></tr><tr><td>W10</td><td>72.1</td><td>66.0</td><td>952.8</td><td>1093.0</td><td>416.8</td><td>457.5</td><td>455.8</td><td>533.5</td><td>536.0</td><td>635.5</td><td>-497.0</td><td>-559.5</td><td>39.0</td><td>76.0</td></tr><tr><td>W11</td><td>72.9</td><td>66.6</td><td>1041.5</td><td>1067.2</td><td>461.5</td><td>432.2</td><td>519.5</td><td>442.2</td><td>580.0</td><td>635.0</td><td>-522.0</td><td>-625.0</td><td>58.0</td><td>10.0</td></tr><tr><td>W12</td><td>71.7</td><td>66.7</td><td>988.0</td><td>1067.8</td><td>446.5</td><td>456.3</td><td>518.0</td><td>543.3</td><td>541.5</td><td>611.5</td><td>-470.0</td><td>-524.5</td><td>71.5</td><td>87.0</td></tr><tr><td>W13</td><td>72.7</td><td>66.9</td><td>1001.5</td><td>1096.6</td><td>436.5</td><td>468.1</td><td>504.5</td><td>562.1</td><td>565.0</td><td>628.5</td><td>-497.0</td><td>-534.5</td><td>68.0</td><td>94.0</td></tr><tr><td>W14</td><td>71.4</td><td>67.2</td><td>991.5</td><td>1148.7</td><td>461.0</td><td>473.7</td><td>527.5</td><td>568.2</td><td>530.5</td><td>675.0</td><td>-464.0</td><td>-580.5</td><td>66.5</td><td>94.5</td></tr><tr><td>W15</td><td>71.0</td><td>65.8</td><td>1045.0</td><td>1164.6</td><td>467.5</td><td>473.6</td><td>538.5</td><td>531.6</td><td>577.5</td><td>691.0</td><td>-506.5</td><td>-633.0</td><td>71.0</td><td>58.0</td></tr><tr><td>W16</td><td>71.6</td><td>66.2</td><td>993.0</td><td>1159.7</td><td>414.0</td><td>478.7</td><td>437.0</td><td>534.2</td><td>579.0</td><td>681.0</td><td>-556.0</td><td>-625.5</td><td>23.0</td><td>55.5</td></tr><tr><td>W17</td><td>71.3</td><td>67.0</td><td>1006.9</td><td>1116.7</td><td>430.9</td><td>450.2</td><td>508.4</td><td>545.7</td><td>576.0</td><td>666.5</td><td>-498.5</td><td>-571.0</td><td>77.5</td><td>95.5</td></tr><tr><td>W18</td><td>70.9</td><td>66.2</td><td>1041.4</td><td>1114.4</td><td>453.4</td><td>496.9</td><td>527.4</td><td>563.4</td><td>588.0</td><td>617.5</td><td>-514.0</td><td>-551.0</td><td>74.0</td><td>66.5</td></tr><tr><td>Min</td><td>70.4</td><td>65.6</td><td>885.8</td><td>1063.4</td><td>233.8</td><td>336.2</td><td>273.3</td><td>368.2</td><td>484.0</td><td>611.5</td><td>-612.5</td><td>-711.0</td><td>23.0</td><td>10.0</td></tr><tr><td>Max</td><td>72.9</td><td>68.0</td><td>1074.1</td><td>1223.4</td><td>494.9</td><td>515.4</td><td>542.3</td><td>596.4</td><td>652.0</td><td>743.0</td><td>-418.5</td><td>-524.5</td><td>84.0</td><td>96.0</td></tr><tr><td>Mean</td><td>71.7</td><td>66.6</td><td>1012.0</td><td>1124.2</td><td>440.0</td><td>459.0</td><td>502.2</td><td>518.4</td><td>572.0</td><td>665.3</td><td>-509.8</td><td>-605.8</td><td>62.2</td><td>59.5</td></tr><tr><td>S. Dev.</td><td>0.38</td><td>0.49</td><td>33.54</td><td>34.33</td><td>24.97</td><td>23.48</td><td>30.23</td><td>28.58</td><td>28.18</td><td>33.65</td><td>30.22</td><td>34.63</td><td>12.53</td><td>13.38</td></tr></tbody></table><table><caption>Statistical significance of the sources of variation (probability > F) from ANOVA</caption><tbody><tr><td>Env</td><td>&lt;.0001</td><td>&lt;.0001</td><td>&lt;.0001</td><td>0.0060</td><td>&lt;.0001</td><td>&lt;.0001</td><td>0.3778</td></tr><tr><td>Clon</td><td>&lt;.0001</td><td>&lt;.0001</td><td>&lt;.0001</td><td>&lt;.0001</td><td>&lt;.0001</td><td>&lt;.0001</td><td>&lt;.0001</td></tr><tr><td>E°C</td><td>0.6549</td><td>0.8442</td><td>0.6336</td><td>0.8300</td><td>0.9567</td><td>0.7993</td><td>0.7949</td></tr></tbody></table>

PT, pasting temperature; PV, peak viscosity; HP, hot paste (through) viscosity; CP, cool paste (end) viscosity; BD, breakdown; SB, setback; CS, consistency. At the bottom the statistical significances for Environment (Env), Genetic (Clon) and Genotype-by-Environment interactions (GxE) effects are presented.

in samples from the IA environment. The average Dx(50) values were 13.98 (±1.05) and 15.54 (±0.89) µm, respectively. These averages were significantly different, as well (p < 0.0001). Wildtype samples from the IA environment ranged from 12.18 to 15.95 µm. Although wildtypes and waxy starch clones are genetically different for many loci (in addition to the GBSS gene) and the number of samples analyzed is limited, there is a clear indication that waxy starches tend to have slightly larger granules.

Results on gel clarity, swelling and RVA pasting behavior tend to confirm the findings of previous works.[5-7] Waxy starches produce gels with higher clarity (66.7% and 66.5% at LA and IA, respectively Table 4) than wildtype starches (61.9% and 62.7%, respectively Table 2). There were significant differences in gel clarity among wildtype and waxy genotypes. However, the other sources of variation had little influence and their F-tests were not significant.

Solubility was markedly higher in wildtype versus waxy starches (Table 2 and 4, respectively). There were no

significant differences among clones, environments and environment-by-clones interaction (GxE) effects in wildtype starches (Table 2). On the other hand significant difference were observed in waxy starches for differences among environments or clones. There was a sharp contrast between swelling for wildtype (average across environments 29.2 g g⁻¹) and waxy starches (59.9 g g⁻¹) and significant variation among clones with wildtype starch. GxE for swelling was not significant for either type of starch.

**Table 3** and 5 present the RVA results. Difference in PT between wildtype and waxy starches was negligible. There was a significant effect of the environment (LA vs IA) on PT in wildtype (71.5 vs. 67.9 °C, Table 3) and waxy (71.7 and 66.6 °C, Table 5) starches. PTs among clones producing waxy starch were significantly different but that was not the case among wildtype clones (Table 5 and 3, respectively). GxE effects were not significant for PT.

One of the most distinctive difference among waxy and wildtype starches is in PV. Wildtype starches showed PV of 653.6 and 858.4 cP in LA and IA environments, respectively (Table 3).
<figure><img src="image_2.png" /><figcaption>**Figure 1.** Illustration showing key amylogram differences among starches from waxy and wildtype genotypes grown at low and intermediate altitudes in the first experiment (left). Starches from mid-altitude show higher peak viscosity compared with those from lower altitudes. Waxy starches, independently of their origin show higher peak viscosity compared with wildtype starches. Amylograms on the right (second experiment) illustrate the differences between intermediate and high altitudes on starch properties of a single waxy cassava genotype. The higher the altitude the higher the peak viscosity (except for Corpoica's sample).</figcaption></figure>

These differences were statistically significant. PV of waxy starches were considerably higher (1012.0 and 1124.2 cP respectively for LA and IA, Table 5). In both starch types, the environment sources of variation showed a significant *F*-test. GxE effects were not significant for PV. Key points in the amylograms from wildtype and waxy starches coming from LA and IA environments were used to draw the hand-made graph presented in **Figure 1**. Differences for PV can be appreciated in this illustration. It is clear that waxy starches have higher PV than wildtypes and also that the IA environment induced higher PV than the LA location.

Two factors may have contributed to differences in PV: first, the higher swelling capacity of waxy starches.[31] Second, the larger granule size of waxy starches may have also played a role in the observed differences in functional properties.

The contrasting results for PV became negligible for HP, which was identical for wildtype and waxy starches from LA (Table 3 and 5) and very similar for IA (461.4 vs. 459.0 cP for wildtype and waxy starches, respectively). The only significant effect for HP in wildtype genotypes was for differences among clones (*p* = 0.0451). In the case of waxy starches both, environment and clones, were significant, whereas GxE was not (Table 3 and 5). The relative values for PV and HP in waxy and wildtype starches result in contrasting results for BD, which was (as expected) considerably larger in the former (572.0 and 665.3 cP, Table 5) than in the latter (213.6 and 397.1 cP, Table 3). For both types of starches, environment and clones had significant *F*-tests, but GxE effects were not significant.

CP was higher for wildtype (598.3 and 680.6 cP, Table 3) than for waxy starches (502.2 and 518.4 cP, Table 5). The formation of

<table><caption>**Table 6.** Correlation coefficients among several parameters quantified in genotypes producing wildtype and waxy starch grown in low and intermediate environments.</caption><thead><tr><th></th><th>PC</th><th>SO</th><th>SW</th><th>PT</th><th>PV</th><th>HP</th><th>CP</th><th>BD</th><th>SB</th><th>CS</th></tr></thead><tbody><tr><th>PC</th><td>1.00</td><td>0.45**</td><td>-0.26**</td><td>-0.05<sup>NS</sup></td><td>0.36**</td><td>-0.15<sup>NS</sup></td><td>-0.31**</td><td>0.45**</td><td>-0.45**</td><td>-0.35**</td></tr><tr><th>SO</th><td></td><td>1.00</td><td>-0.89**</td><td>-0.13<sup>NS</sup></td><td>0.80**</td><td>0.11<sup>NS</sup></td><td>-0.60**</td><td>0.85**</td><td>-0.92**</td><td>-0.82**</td></tr><tr><th>SW</th><td></td><td></td><td>1.00</td><td>0.14<sup>NS</sup></td><td>-0.82**</td><td>-0.21**</td><td>0.52**</td><td>-0.84**</td><td>0.90**</td><td>0.78**</td></tr><tr><th>PT</th><td></td><td></td><td></td><td>1.00</td><td>-0.50**</td><td>-0.19*</td><td>-0.15<sup>NS</sup></td><td>-0.46**</td><td>0.33**</td><td>-0.06<sup>NS</sup></td></tr><tr><th>PV</th><td></td><td></td><td></td><td></td><td>1.00</td><td>0.36</td><td>-0.09<sup>NS</sup></td><td>0.92**</td><td>-0.86**</td><td>-0.49**</td></tr><tr><th>HP</th><td></td><td></td><td></td><td></td><td></td><td>1.00</td><td>0.79**</td><td>-0.03<sup>NS</sup></td><td>0.11<sup>NS</sup></td><td>0.29**</td></tr><tr><th>CP</th><td></td><td></td><td></td><td></td><td></td><td></td><td>1.00</td><td>-0.42**</td><td>0.58**</td><td>0.81**</td></tr><tr><th>BD</th><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>1.00</td><td>-0.97**</td><td>-0.64**</td></tr><tr><th>SB</th><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>1.00</td><td>0.81**</td></tr><tr><th>CS</th><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>1.00</td></tr></tbody></table>

PC, paste clarity; SO, solubility; SW, swelling power; PT, pasting temperature; PV, peak viscosity; HP, hot paste (through) viscosity; CP, cool paste (end) viscosity; BD, breakdown; SB, setback; CS, consistency.

NS, *, **: non significant, significant at 5% and significant at 1% probability levels, respectively.
<table><caption>**Table 7.** Starch quality traits from waxy clone AM 206-5 grown in five different environments in Colombia.</caption><thead><tr><th>Location</th><th>Altitude [m.a.s.l.]</th><th>PC [%]</th><th>SO [% db]</th><th>SW [g g<sup>-1</sup>]</th><th>PT [°C]</th><th>PV [cP]</th><th>HP [cP]</th><th>CP [cP]</th><th>BD [cP]</th><th>SB [cP]</th><th>CS [cP]</th></tr></thead><tbody><tr><td>Sonso</td><td>934</td><td>57.6</td><td>13.4</td><td>56.3</td><td>64.0</td><td>1023.0</td><td>673.5</td><td>702.5</td><td>349.5</td><td>-320.5</td><td>29.0</td></tr><tr><td>CIAT</td><td>965</td><td>61.5</td><td>9.0</td><td>59.6</td><td>66.8</td><td>1054.0</td><td>454.5</td><td>522.5</td><td>599.5</td><td>-531.5</td><td>68.0</td></tr><tr><td>Corpoica</td><td>1001</td><td>65.2</td><td>10.9</td><td>65.9</td><td>66.7</td><td>968.5</td><td>465.0</td><td>545.5</td><td>503.5</td><td>-423.0</td><td>80.5</td></tr><tr><td>Quilichao</td><td>1071</td><td>58.5</td><td>11.5</td><td>59.2</td><td>66.1</td><td>1163.0</td><td>509.0</td><td>555.5</td><td>654.0</td><td>-607.5</td><td>46.5</td></tr><tr><td>Darien</td><td>1485</td><td>59.2</td><td>6.9</td><td>54.4</td><td>62.9</td><td>1437.0</td><td>594.5</td><td>637.0</td><td>842.5</td><td>-800.0</td><td>42.5</td></tr><tr><td>LSD (5%)</td><td></td><td>1.37</td><td>1.28</td><td>1.07</td><td>0.27</td><td>11.07</td><td>18.88</td><td>25.51</td><td>17.65</td><td>23.02</td><td>18.66</td></tr></tbody></table>

PC, paste clarity; SO, solubility; SW, swelling power; PT, pasting temperature; PV, peak viscosity; HP, hot paste (through) viscosity; CP, cool paste (end) viscosity; BD, breakdown; SB, setback; CS, consistency.

a network of retrograded amylose helices during cooling is the most likely explanation factor for this observation.[32,33] CP tended to be higher in starches from plants growing at IA. GxE effects for CP were not significant. Environment resulted in significant F-tests for both types of starches (Table 3 and 5) but variations among clones was only significant in waxy clones (Table 5).

The relatively low CP in cassava starches resulted in negative values for SB, particularly for waxy genotypes (–509.8 and –605.8 cP, Table 5). In the case of wildtype starches (Table 3) environment and genetic effects were significant, but that was not the case for their interaction. The same conclusions can be drawn for waxy starches (Table 5).

Finally, CS was smaller in waxy (62.2 and 59.5 cP, Table 5) than in wildtype starches (158.3 and 219.3 cP, Table 3), due to the retrogradation of amylose during cooling. Environment had a strong and significant effect among wildtypes, whereas the effect on waxy clones was negligible. GxE effects were not significant.

**Table 6** presents the correlation coefficients among the different variables evaluated taking into consideration data from both locations. Correlations among the same variables but for LA or IA environments were also measured (data not presented). In general correlations across environment, at LA or IA were similar, except for those involving PT. The solubility and swelling power (SO and SW) influence pasting properties, in particular peak viscosity (PV).

## 3.2. Second Experiment

The effect of environment on starch quality of a single waxy genotype (AM 206–5) grown in five locations is presented in **Table 7**. The location effect in this experiment was highly significant for all variables. The main objective of this trial was to assess the impact of HA on starch functional properties but only on a single waxy starch genotype. Peak viscosity is positively associated with altitude (Table 7 and Figure 1), further supporting the results obtained in the first experiment. The molecular mechanisms behind the increase in PV with altitude are not clear yet.

This experiment was actually conducted a decade ago but the results were not published because of the limited information provided by it. However, it complements nicely the results from the first experiment and thus its inclusion in the present article.

Unfortunately, these old starch samples were no longer available for further analysis of granule size and other structural properties.

## 4. Conclusions

GxE effects on the properties of waxy and wildtype cassava starches were not significant, whereas environment and/or genotypes within starch type sources of variation were generally significant (except for PC and SO). Results from the two experiments cannot be compared because different genotypes and environments were used. From both experiments it is clear, however, that PV was positively correlated with altitude. In the first experiment PVs from wildtype genotypes were 653.6 and 858.4 cP, respectively, for LA and IA. Similar results were observed in waxy genotypes (1012.0 and 1124.2 cP for LA and IA, respectively). In the second experiment, waxy starches from plants grown at the highest altitude (Darien, 1485 m) showed the highest PV (1437 cP), followed by those at the second highest altitude (Quilichao, 1071 m) with 1163 cP. The remaining locations, with altitudes ranging from 934 to 1001 m, had an average PV of 1052 cP.

Many root and tuber crops can be grown in a wide array of altitudes (e.g., from sea level up to 1800 m.a.s.l, in the case of cassava) in the tropics. Results from this study suggest that it is relevant to pay attention to the altitude at which crops are grown when reporting starch functional properties. Perhaps temperature is the most distinctive difference between the environments used in this study. We can hypothesize that variation in temperature differentially influenced the activity of enzymes involved in starch biosynthesis (in agreement with Buléon et al.[34]).

As reported in the literature,[5–7] PV was higher in waxy starch genotypes than wildtypes, irrespective on the environment where plants were grown. CP, on the other hand, tended to be higher in wildtype starches, also irrespective of the environment where plants were grown (Figure 1). Differences for HP, although statistically significant were small, with no clear trend between waxy and wildtype starches (Figure 1).

Waxy starches grown in LA tended to have smaller granules than those from IA. The difference in granule sizes may have indirectly affected the functional properties of the starches, in particular PV. Similarly, the waxy starch trait seemed to be related to slightly larger granule size compared with wildtype samples. Further research, with starch samples from different genotypes, need to confirm this preliminary finding.
## Abbreviations

BD, breakdown; CP, cool paste (end) viscosity; CS, consistency; GxE, genotype-by-environment interaction; HP, hot paste (through) viscosity; HA, high altitude; IA, Intermediate altitude; LA, low altitude; PC, Paste clarity; PT, pasting temperature; PV, peak viscosity; SB, setback; SO, solubility; SW, swelling power.

## Acknowledgments

This work was conducted with financial support from the following sources: CGIAR Research Program on Roots, Tubers and Bananas (RTB) and Ingredion Colombia S.A.

## Conflict of Interest

The authors declare no conflict of interest.

## Keywords

correlations, root and tuber crops, stability, tropical sources of starch, waxy starch

Received: October 16, 2017

Revised: April 6, 2018

Published online: July 27, 2018

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