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標題: | 黑棘蟻聚落分工機制及生物時鐘表現 Models of Division of Labor and Expression of Biological Clock in the Colony of Gray-Black Spiny Ant, Polyrhachis dives Smith (Hymenoptera: Formicidae) |
作者: | Jaw-Jinn Jong 鍾兆晉 |
指導教授: | 李後晶(How-Jing Lee) |
關鍵字: | 分工,生物時鐘,黑棘蟻, division of labor,biological clock,Polyrhachis dives, |
出版年 : | 2008 |
學位: | 博士 |
摘要: | 典型的社會性昆蟲的聚落,是自我組織的單位,是去中心化的系統。聚落整體的表現是由所有個體個別的行為所匯聚,是一系列獨立活動的串接。分工是所有研究社會性昆蟲聚落行為方面最基礎、最廣泛的研究領域,分工的研究乃探討包括個別工蟻行為的整合、進入聚落階層後工作的分配,以及如何調節分工以促進聚落的效能等項目。
本論文是以「分工對社會性昆蟲行為適應之意義」為主軸,探討分工在無形態多型性的社會結構下發展出什麼模式,以應對環境的變化。具體的研究方法是選擇黑棘蟻為研究對象,確定此種螞蟻的社會組織、生殖階級的變化以及工蟻亞階級的調控。黑棘蟻的聚落組成可分為缺后、單后及多后三種,聚落的循環則是分封創設與獨立創設同時存在,生殖階級的複雜行為模式,是在蟻類中罕見的,包括處女蟻后、蟻后及雄蟻,都會單次交尾與多次交尾,也會雜交與自交。最特別的是,處女蟻后在一年內有兩次出現的高峰,而雄蟻則終年都可發現。根據野外的長期監測,黑棘蟻可以形成獨立聚落,也容易形成超級聚落,而超級聚落內不同聚落依功能及組成來分,可區別出四類聚落,包括:中心聚落、生殖聚落、支持聚落以及保留聚落,顯示超級聚落的維持依賴不同功能聚落的分工。比較不同社會組織的聚落在生殖季和非生殖季的差別,發現包括聚落大小、性別比例以及雄蟻先成的情形都有顯著的差異。 焦點轉移到巢內,黑棘蟻的工蟻亞階級則以行為譜分析被區分為內勤工蟻、警衛蟻以及外勤工蟻,另外再加上初生工蟻,這四群工蟻在巢內的行為包括:休息的比例、休息的長度、交哺、接觸頻率以及自動甦醒的機率,這些行為都有顯著差異,但是在形態學和解剖學上的比較,則四群工蟻都沒有顯著的不同。研究顯示在巢內有大部分的螞蟻都在休息,其實休息是任務轉換的重要步驟,休息後不同工蟻亞階級可能會互換角色,而如此的結果也證實了本種螞蟻之工蟻亞階級的分工是呈現較弱的年齡行為的多型性,以及較有彈性的任務分配。 既然休息是轉換任務的關鍵因素,那麼黑棘蟻蟻巢內間歇性的全面活動則是使整個聚落同時甦醒的重要現象。這種間歇性的全面活動在有后聚落中竟然呈現自由律動,顯示螞蟻聚落的超有機體存在類似脊椎動物個體的脈搏,這個有自由律動的間歇性全面活動,則會在缺后聚落中失去規律。 至於職務的分配,研究結果顯示:大型聚落、小型聚落和缺后聚落之間差異皆不明顯,顯示螞蟻社會為了要維持生存必須先調整分工比例,過度的角色偏斜都會使聚落組織失去平衡而喪失競爭力。在黑棘蟻聚落成員中,比較特殊的、有別於其他螞蟻的是警衛蟻這個角色,研究發現警衛蟻的數目會隨著聚落大小、食物量和食物密度改變而改變,而覓食歷程中外勤工蟻交互出現的時間序列也被完整的分析出來。 在實驗室中,黑棘蟻個別的個體被連續的紀錄儀器完整紀錄活動量和活動時間,發現黑棘蟻的覓食和生殖行為是日行性。除了產卵蟻后和內勤工蟻之外,所有的螞蟻階級都表現出日週律動。因為隔離的螞蟻有可能會改變角色,所以設計生物檢定法去測量實驗結束後工蟻的角色轉換。研究顯示不同階級和不同亞階級都會表現日週律動,但是產卵和育幼則會產生掩蓋效應,這些現象都是由超有機體的概念進行解釋與探討。 Typical social structure of ant is self-organized and decentralized system. The whole colony displays assembled behaviors which are composed independently by individuals' specific behavior. Division of labor, the most basic and extensive research field of studying ant, is the backbone to assemble individual-mates behaviors. Many researches attempt to probe into task partition of individual workers in the whole colony and how to regulate division of labor in order to promote colony’s efficiency. This thesis focused on the adaptation significance of division of labor in the gray-black spiny ant, Polyrhachis dives. I attempt to understand its social structure, the regulation of sexual and non-sexual castes, and task-shift of worker subcastes. Three types of colony as queenless, monogyny and polygyny were found in nature condition. The colony funding could be either by budding or independent which could be found at the same time in the same habitat. In the nature colony, several reproductive castes, including virgin queens, queens and males, were commonly found and could perform either monoandrous (monogamous) or polyandrous (polygamous) which would result in interchange of out-breeding and in-breeding. Although virgin queens appeared twice within one year, males could be found all year long. Based on my long-term observation, P. dives could form not only independent colony, but also super-colony. The uniqueness of the super-colony of P. dives was its 4 distinguish function and structure colonies such as central colony, reproductive colony, supportive colony and reserve colony. The division of labor in super-colony also relied on temporal regulation of functions among 4 colonies. Seasonal variation was reflected on colony size, sex ratio and presence of male caste. Under detailed observation, ethogram of nurse, guard, forager and callow worker was constructed to reveal all the behavior repertoires in the nest. Although there was no significant difference on morphological characteristics among four workers, there were significant difference on resting proportion, resting duration, trophallaxis, encounter rate with nest mates, and spontaneous activity ratio. Special attention was on resting because it was the most common behavior in the nest and served as an essential stage for task shifting. Different sub-castes of worker might have different rates of task shifting which represented weak age-polyethism and flexible task partition. Since resting was a key factor of changing tasks, pulses of activity following resting to revive the whole colony was the important phenomenon to communicate with nest-mates for division of labor. The daily pulses of activity expressing free running rhythm in queenright colonies under DD condition, revealed pulse-like phenomenon in heart beat of vertebrate which was high at active phase and low at inactive phase. However, this circadian rhythm was disappeared in queenless colonies. In the study of task partition in large, small, and queenless colony, the proportion of tasks was reflecting the efficient proportion for survive of the colony. Artificial biased the proportion would trigger the task shifting to balance out the excessive tasks. Guard was the unique task found in P. dives. The recruitment of guard was depended on colony size, food quality and food density. Temporal analysis on the existence of foragers and guards at food source revealed the dynamics of task partitioning between these sub-castes. Continuous monitoring of the locomotion of the gray-black spiny ant, Polyrhachis dives, showed a diurnal nature in foraging and reproduction. Furthermore, all the castes except dominant queens and nurses expressed circadian locomotor rhythms. Since isolation could cause task-shifting in worker castes, a series of behavioral bioassays were designed to determine the task of workers at the end of locomotor detection. The differential expression of circadian locomotor rhythms among castes reflected a masking effect on locomotion by social status or task demands. This unique phenomenon is discussed in the context of the “super-organism” concept. |
URI: | http://tdr.lib.ntu.edu.tw/jspui/handle/123456789/40856 |
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